Facilitate Invasion Success Research Articles

Bold-shy personality traits of globally invasive, native and hatchery-reared fish.

Bold behaviour of non-native species is hypothesized to facilitate invasion success, yet extreme boldness in wild and domesticated animals can be maladaptive. The purpose of this study was to compare individual behaviour among Australian native hatchery-reared (n = 33) and wild (n = 38) Murray cod (Maccullochella peelii) with invasive common carp (Cyprinus carpio; n = 30). Three laboratory tests measured individual behaviour: (1) emergence from a shelter, (2) exploration of a novel environment, and (3) approaching a predator. Wild invasive carp and hatchery-reared cod were generally faster and more likely to emerge and explore novel environments when compared with wild Murray cod. The 'bold-type' behaviours of hatchery-reared native cod were more like invasive carp than they were to 'shy-type' wild conspecifics, yet an important difference was that hatchery-reared cod spent substantially more time near a large predator while carp rapidly escaped. We suggest that these results are consistent with a bold-type invasion syndrome in invasive carp and learned boldness of hatchery-reared Murray cod. The propensity of invasive carp to rapidly explore and enter new environments, along with a fast predator escape response may have been important to their invasion success, while extreme risk-taking and predator naivety of hatchery-reared Murray cod may exacerbate post-release mortality rates in fisheries and conservation stocking programmes.

Quantifying the impacts of symbiotic interactions between two invasive species: the tawny crazy ant (Nylanderia fulva) tending the sorghum aphid (Melanaphis sorghi).

The establishment of new symbiotic interactions between introduced species may facilitate invasion success. For instance, tawny crazy ant (Nylanderia fulva Mayr) is known to be an opportunistic tender of honeydew producing insects and this ants' symbiotic interactions have exacerbated agricultural damage in some invaded regions of the world. The invasive sorghum aphid (Melanaphis sorghi Theobald) was first reported as a pest in the continental United States-in Texas and Louisiana-as recent as 2013, and tawny crazy ant (TCA) was reported in Texas in the early 2000s. Although these introductions are relatively recent, TCA workers tend sorghum aphids in field and greenhouse settings. This study quantified the tending duration of TCA workers to sorghum aphids and the impact of TCA tending on aphid biomass. For this study aphids were collected from three different host plant species (i.e., sugarcane, Johnson grass, and sorghum) and clone colonies were established. Sorghum is the main economic crop in which these aphids occur, hence we focused our study on the potential impacts of interactions on sorghum. Quantification of invasive ant-aphid interactions, on either stems or leaves of sorghum plants, were conducted in greenhouse conditions. Our results show that although these two invasive insect species do not have a long coevolutionary history, TCA developed a tending interaction with sorghum aphid, and aphids were observed excreting honeydew after being antennated by TCA workers. Interestingly, this relatively recent symbiotic interaction significantly increased overall aphid biomass for aphids that were positioned on stems and collected from Johnson grass. It is recommended to continue monitoring the interaction between TCA and sorghum aphid in field conditions due to its potential to increase aphid populations and sorghum plant damage.

A Behavioral Syndrome Linking Boldness and Flexibility Facilitates Invasion Success in Sticklebacks.

AbstractFor a species to expand its range, it needs to be good at dispersing and also capable of exploiting resources and adapting to different environments. Therefore, behavioral and cognitive traits could play key roles in facilitating invasion success. Marine threespined sticklebacks (Gasterosteus aculeatus) have repeatedly colonized freshwater environments and rapidly adapted to them. Here, by comparing the behavior of hundreds of lab-reared sticklebacks from six different populations, we show that marine sticklebacks are bold, while sticklebacks that have become established in freshwater lakes are flexible. Moreover, boldness and flexibility are negatively correlated with one another at the individual, family, and population levels. These results support the hypothesis that boldness is favored in invaders during the initial dispersal stage, while flexibility is favored in recent immigrants during the establishment stage, and they suggest that the link between boldness and flexibility facilitates success during both the dispersal stage and the establishment stage. This study adds to the growing body of work showing the importance of behavioral correlations in facilitating colonization success in sticklebacks and other organisms.

The potential impacts of invasions on native symbionts

Symbionts, including parasites, pathogens, and mutualists, can play important roles in determining whether or not invasions by host species will be successful. Loss of enemies from the native habitat, such as parasites and pathogens, can allow for higher invader fitness in the invaded habitat. The presence of mutualists (e.g., pollinators, seed dispersers, mycorrhizae, and rhizobial bacteria) in the invaded habitat can facilitate invasion success. Although there has been a great deal of research focusing on how invading hosts may benefit from enemy losses or mutualist gains, far less attention has focused on how native symbiont populations and communities respond to invasion by non‐indigenous hosts and symbionts. In this paper, we present a conceptual framework examining how symbionts such as parasites, pathogens, commensals, and mutualists can influence invader success and whether these native symbionts will benefit or decline during invasion. The first major factor in this framework is the competence of the invading host relative to the native hosts. Low‐ or non‐competent hosts that support few if any native symbionts could cause declines in native symbiont taxa. Competent invading hosts could potentially support native parasites, pathogens, commensals, and mutualists, especially if there is a closely related or similar host in the invaded range. These symbionts could inhibit or facilitate invasion or have no discernible effect on the invading host. An understanding of how native symbionts interact with competent versus non‐competent invading hosts as well as various invading symbionts is critical to our understanding of invasion success, its consequences for invaded communities and how native symbionts in these communities will fare in the face of invasion.

Hybridization between two introduced, invasive crayfish species in the upper Juniata River system, Pennsylvania, USA

Abstract Hybridization of non-native crayfishes with native crayfishes can facilitate invasion success of the non-native. We report novel genetic evidence of hybridization between rusty crayfish, Faxonius rusticus (Girard, 1852), and Allegheny crayfish, F. obscurus (Hagen, 1870), two non-native species in the upper Juniata River basin of central Pennsylvania. We used double-digest restriction-associated DNA sequencing, which yielded 28,031 single nucleotide polymorphisms, to characterize the extent of hybridization. Two of eight individuals with mixed character traits were identified as the progeny of hybrids backcrossed into rusty crayfish. One individual, presumed to be a rusty crayfish in the field, was genetically identified as a hybrid backcross to rusty crayfish. The three crayfish taxa (rusty, Allegheny, and intermediates) are virtually indistinguishable based on typical morphological characters. Key character traits that distinguish rusty and Allegheny crayfishes can be mixed atypically in certain individuals (i.e., intermediates). Our results indicate that introgressive hybridization has occurred between rusty and Allegheny crayfishes, and that at least some of the character-trait intermediate individuals have hybrid ancestry. Hybridization may be an additional means by which invasive rusty crayfish increase their fitness and dominance at the expense of existing species and their habitat.

Genome-Wide SNP Analysis Reveals Multiple Paternity in Burmese Pythons Invasive to the Greater Florida Everglades

Reproductive strategies are an essential component of invasion ecology that influence invasion success and rates of population growth. Burmese Pythons (Python bivittatus) are large constrictor snakes that were introduced to the Greater Everglades Ecosystem of southern Florida, USA, from Asia. Since their introduction, these giant constrictors have spread throughout wetlands of southern Florida while increasing in abundance and causing declines in the native species upon which they prey. Multiple paternity in reproduction could facilitate invasion success by increasing the genetic diversity produced within each reproductive event. We used Diversity Arrays Technology genome-wide genotyping to assess multiple paternity in the progeny of wild Burmese Pythons in Florida. We analyzed >4,000 single nucleotide polymorphisms from 153 neonates belonging to 4 clutches collected in southwestern Florida. Complementary hierarchical and K-means clustering analyses of the genetic distances within clutches revealed that three clutches were each fertilized by two sires, with a fourth fertilized by a single sire. The proportions of offspring attributable to each sire within multiple paternity clutches ranged from nearly even to highly skewed. Analysis of multivariate dispersion showed significantly increased genetic variability in the multiple paternity clutches. These results improve our understanding of the reproductive strategy and invasion potential of a giant constrictor with significant ecological impacts.

Reproductive Strategies That May Facilitate Invasion Success: Evidence From a Spider Mite.

With significant surge of international trade in recent decades, increasingly more arthropod species have become established outside their natural range of distribution, causing enormous damage in their novel habitats. However, whether a species can colonize its new environment depends on its ability to overcome various barriers which may result in establishment failure, such as inbreeding depression and difficulty to find mates. Here, we used a haplodiploid pest, Tetranychus ludeni Zacher (Acari: Tetranychidae), which is native to Europe but now cosmopolitan, to investigate whether its reproductive strategies have facilitated its invasion success, providing knowledge to develop programs for prediction and management of biological invasions. We show that inbreeding had no negative influence on female reproductive outputs and longevity over 11 successive generations, allowing mother-son and brother-sister mating to occur at the invasion front without adverse consequences in fitness. Virgin females produced maximum number of sons in their early life to ensure subsequent mother-son mating but later saved resources to prolong longevity for potential future mating. Females maximized their resource allocation to egg production immediately after mating to secure production of maximum number of both daughters and sons as early as possible. Furthermore, mated females with mating delay increased proportion of daughters in offspring produced to compensate the loss of production of daughters during their virgin life. We suggest that the lack of inbreeding depression in successive generations and the ability to adjust resource allocations depending whether and when mating occurs may be the key features that have facilitated its invasion success.

Phylogenomics resolves the invasion history of Acacia auriculiformis in Florida

AbstractAimUnderstanding the genetic structure of plants in their native range is crucial when reconstructing the invasion history of weeds. This information allows researchers to pinpoint the provenance of invasive plants and to test the importance of genetic admixture in facilitating invasion success. We assessed genetic structuring across the native range of A. auriculiformis, to determine whether genetic admixture contributes to the success of this weed in its introduced range, and test for rapid adaptation to environmental conditions in the invasive lineage.LocationAustralia, Papua New Guinea, Florida.TaxonEarleaf acacia.MethodsWe sampled A. auriculiformis from across its entire native distribution (northern Australia, Papua New Guinea) and its invasive range in Florida, and used genotyping by sequencing (GBS) to assess population structuring.ResultsPrincipal component analysis, based on 9591 single‐nucleotide polymorphisms, indicated significant differentiation among samples from Papua New Guinea, the Northern Territory (Australia) and north Queensland (Australia). Florida samples also formed a distinct cluster, with these samples most closely related to samples from the Northern Territory. These results indicate that the Florida A. auriculiformis lineage most likely originates from the Northern Territory, with no evidence that plants were introduced from different parts of the native range. We found evidence of allelic shifts in the Florida population, suggesting rapid adaptation to environmental conditions may contribute the success of the invasive lineage.Main conclusionsTwo well‐known biogeographic barriers—the Carpentaria Gap and Torres Strait—have caused genome‐wide divergence among A. auriculiformis plants from north Queensland, Northern Territory and Papua New Guinea. The taxonomic status of these allopatric populations should be further assessed. As the Florida lineage originated in the Northern Territory, the search for potential biological control agents should be focused in this region. Our results also demonstrate how artificial selection and strong genetic drift may cause introduced plants to have a unique genetic make‐up not found in the native range.

The role of arbuscular mycorrhizal fungi in nonnative plant invasion along mountain roads.

Plant associated mutualists can mediate invasion success by affecting the ecological niche of nonnative plant species. Anthropogenic disturbance is also key in facilitating invasion success through changes in biotic and abiotic conditions, but the combined effect of these two factors in natural environments is understudied. To better understand this interaction, we investigated how disturbance and its interaction with mycorrhizas could impact range dynamics of nonnative plant species in the mountains of Norway. Therefore, we studied the root colonisation and community composition of arbuscular mycorrhizal (AM) fungi in disturbed vs undisturbed plots along mountain roads. We found that roadside disturbance strongly increases fungal diversity and richness while also promoting AM fungal root colonisation in an otherwise ecto-mycorrhiza and ericoid-mycorrhiza dominated environment. Surprisingly, AM fungi associating with nonnative plant species were present across the whole elevation gradient, even above the highest elevational limit of nonnative plants, indicating that mycorrhizal fungi are not currently limiting the upward movement of nonnative plants. We conclude that roadside disturbance has a positive effect on AM fungal colonisation and richness, possibly supporting the spread of nonnative plants, but that there is no absolute limitation of belowground mutualists, even at high elevation.

Chemical responses of an invasive plant to herbivory and abiotic environments reveal a novel invasion mechanism

Invasive plant environments differ along latitudes and between native and introduced ranges. In response to herbivory and abiotic stresses that vary with latitudes and between ranges, invasive plants may shift their secondary chemicals to facilitate invasion success. However, it remains unclear whether and how invasive plant chemical responses to herbivory and chemical responses to abiotic environments are associated. We conducted large scale field surveys of herbivory on the invasive tallow tree (Triadica sebifera) along latitudes in both its native (China) and introduced ranges (United States) and collected leaf samples for analyses of tannins and flavonoids. We used data on climate and solar radiation to examine these chemical responses to abiotic environments and their variations along these latitudes and between ranges. We also re-analyzed previously published data from multiple common garden experiments on tallow tree to investigate genetic divergence of secondary chemical concentrations between introduced and native populations. We found foliar tannins and herbivory (chewing, sucking) were higher in the native range compared to the invasive range. Allocation to tannins versus flavonoids decreased with latitude in the native range but did not vary in the invasive range. Analyses of previously published common garden experimental data indicated genetic divergence contributes to chemical concentration differences between ranges. Our field data further indicated that the latitudinal patterns were primarily phenotypic responses to herbivory in China while in US they were primarily phenotypic responses to abiotic environments. The variation of tannins may be linked to flavonoids, given tannins and flavonoids share a biosynthesis pathway. Together, our results suggest that invasive plants adjust their secondary metabolism to decrease chemicals that primarily defend against herbivory and increase those that help them to respond to their abiotic environment. These findings deepen our understanding of how invasive plants adapt to biogeographically heterogeneous environments through trade-offs between secondary chemical responses.

Differences in seed properties and germination between native and introduced populations of Triadica sebifera

Abstract Aims Seeds of many invasive plants germinate more quickly than those of native species, likely facilitating invasion success. Assessing the germination traits and seed properties simultaneously for introduced and native populations of an invasive plant is essential to understanding biological invasions. Here, we used Triadica sebifera as a study organism to examine intraspecific differences in seed germination together with seed characteristics. Methods We measured physical (volume, mass, coat hardness and coat thickness of seeds) and chemical (crude fat, soluble protein, sugar, gibberellins [GA] and abscisic acid [ABA] of kernels) properties of T. sebifera seeds collected in 2017 from 12 introduced (United States) populations and 12 native (China) populations and tested their germination rates and timing in a greenhouse experiment in China. Furthermore, we conducted an extra experiment in the United States using seeds collected in 2016 and 2017 to compare the effects of study sites (China vs. United States) and seed collection time (2016 vs. 2017) on seed germination. Important Findings Seeds from the introduced range germinated faster than those from the native range. Physical and chemical measurements showed that seeds from the introduced range were larger, had higher GA concentrations and GA:ABA ratio, but lower crude fat concentrations compared to those from the native range. There were no significant differences in seed mass, coat hardness and coat thickness or kernel ABA, soluble protein or sugar concentrations between seeds from introduced vs. native ranges. Germination rates were correlated between United States and China greenhouses but germination rates for populations varied between collection years. Our results suggest that larger seeds and higher GA likely contribute to faster germination, potentially facilitating T. sebifera invasion in the introduced range.

Biological invasion: The influence of the hidden side of the (epi)genome

Abstract Understanding the mechanisms underlying biological invasions and rapid adaptation to global change remains a fundamental challenge, particularly in small populations lacking in genetic variation. Two understudied mechanisms that could facilitate adaptive evolution and adaptive plasticity are the increased genetic variation due to transposable elements (TEs), and associated or independent modification of gene expression through epigenetic changes. Here, we focus on the potential role of these genetic and non‐genetic mechanisms for facilitating invasion success. Because novel or stressful environments are known to induce both epigenetic changes and TE activity, these mechanisms may play an underappreciated role in generating phenotypic and genetic variation for selection to act on. We review how these mechanisms operate, the evidence for how they respond to novel or stressful environments, and how these mechanisms can contribute to the success of biological invasions by facilitating adaptive evolution and phenotypic plasticity. Because genetic and phenotypic variations due to TEs and epigenetic changes are often well regulated or “hidden” in the native environment, the independent and combined contribution of these mechanisms may only become important when populations colonize novel environments. A focus on the mechanisms that generate and control the expression of this variation in new environments may provide insights into biological invasions that would otherwise not be obvious. Global changes and human activities impact on ecosystems and allow new opportunities for biological invasions. Invasive species succeed by adapting rapidly to new environments. The degree to which rapid responses to environmental change could be mediated by the epigenome—the regulatory system that integrates how environmental and genomic variation jointly shape phenotypic variation—requires greater attention if we want to understand the mechanisms by which populations successfully colonize and adapt to new environments. A free Plain Language Summary can be found within the Supporting Information of this article.

Modelling the invasion history of Sinanodonta woodiana in Europe: Tracking the routes of a sedentary aquatic invader with mobile parasitic larvae.

Understanding the invasive potential of species outside their native range is one of the most pressing questions in applied evolutionary and ecological research. Admixture of genotypes of invasive species from multiple sources has been implicated in successful invasions, by generating novel genetic combinations that facilitate rapid adaptation to new environments. Alternatively, adaptive evolution on standing genetic variation, exposed by phenotypic plasticity and selected by genetic accommodation, can facilitate invasion success. We investigated the population genetic structure of an Asian freshwater mussel with a parasitic dispersal stage, Sinanodonta woodiana, which has been present in Europe since 1979 but which has expanded rapidly in the last decade. Data from a mitochondrial marker and nuclear microsatellites have suggested that all European populations of S. woodiana originate from the River Yangtze basin in China. Only a single haplotype was detected in Europe, in contrast to substantial mitochondrial diversity in native Asian populations. Analysis of microsatellite markers indicated intensive gene flow and confirmed a lower genetic diversity of European populations compared to those from the Yangtze basin, though that difference was not large. Using an Approximate Bayesian Modelling approach, we identified two areas as the probable source of the spread of S. woodiana in Europe, which matched historical records for its establishment. Their populations originated from a single colonization event. Our data do not support alternative explanations for the rapid recent spread of S. woodiana; recent arrival of a novel (cold‐tolerant) genotype or continuous propagule pressure. Instead, in situ adaptation, facilitated by repeated admixture, appears to drive the ongoing expansion of S. woodiana. We discuss management consequences of our results.

The Braveheart amphipod: a review of responses of invasiveDikerogammarus villosusto predation signals

Predator pressure is a fundamental force driving changes at all levels of the community structure. It may protect native ecosystems from alien species. Therefore, resistance to diverse predators resulting from a universal anti-predator strategy seems crucial for invasion success. We present a comprehensive review of the responses of an invasive amphipod Dikerogammarus villosus to sympatric and allopatric predator signals. We summarize diverse aspects of the gammarid anti-predator strategy, including predator identification, morphological and behavioural adaptations, effectiveness of shelter use and resistance to indirect predator effects. The response of D. villosus is independent of predator species (including totally allopatric taxa), which assures the high flexibility of its predator recognition system. It has a harder exoskeleton and better capability of utilizing shelters compared to other gammarids, resulting in relatively high resistance to predators. Therefore, it can use predator kairomones as indirect food signals (sharing the diet with the predator) and follow the predator scent. This resistance may allow D. villosus to reduce the costs of its physiological responses to predators and sustain growth in their presence. This might facilitate invasion success by increasing its competitive advantage.

Differences in absconding between African and European honeybee subspecies facilitate invasion success of small hive beetles

Small hive beetle (SHB) is an invasive species in populations of European honeybee subspecies, but underlying reasons for SHB success are not well understood. African and European honeybee, Apis mellifera, subspecies differ in absconding, and small hive beetle, greater wax moth (GWM) and ants all can exploit abandoned nests. However, the impact of host absconding on SHB reproduction and the role of GWM and ants as competitors are not known. Here, we conducted a survey in South Africa, Australia and the USA to evaluate SHB and GWM reproduction and foraging by ants in abandoned honeybee colonies. While the impact of competing ants and GWM was not significant, the data show higher SHB reproduction in abandoned nests of European honeybees compared to African ones, but less for GWM. The positive correlation between abandoned protein sources (brood, pollen) on SHB reproduction suggests that the less efficient preparation for absconding by European honeybee subspecies combined with their large colony sizes is a key factor for the invasion success of SHB.

Combining niche shift and population genetic analyses predicts rapid phenotypic evolution during invasion.

The rapid evolution of non‐native species can facilitate invasion success, but recent reviews indicate that such microevolution rarely yields expansion of the climatic niche in the introduced habitats. However, because some invasions originate from a geographically restricted portion of the native species range and its climatic niche, it is possible that the frequency, direction, and magnitude of phenotypic evolution during invasion have been underestimated. We explored the utility of niche shift analyses in the red seaweed Gracilaria vermiculophylla, which expanded its range from the northeastern coastline of Japan to North America, Europe, and northwestern Africa within the last 100 years. A genetically informed climatic niche shift analysis indicates that native source populations occur in colder and highly seasonal habitats, while most non‐native populations typically occur in warmer, less seasonal habitats. This climatic niche expansion predicts that non‐native populations evolved greater tolerance for elevated heat conditions relative to native source populations. We assayed 935 field‐collected and 325 common‐garden thalli from 40 locations, and as predicted, non‐native populations had greater tolerance for ecologically relevant extreme heat (40°C) than did Japanese source populations. Non‐native populations also had greater tolerance for cold and low‐salinity stresses relative to source populations. The importance of local adaptation to warm temperatures during invasion was reinforced by evolution of parallel clines: Populations from warmer, lower‐latitude estuaries had greater heat tolerance than did populations from colder, higher‐latitude estuaries in both Japan and eastern North America. We conclude that rapid evolution plays an important role in facilitating the invasion success of this and perhaps other non‐native marine species. Genetically informed ecological niche analyses readily generate clear predictions of phenotypic shifts during invasions and may help to resolve debate over the frequency of niche conservatism versus rapid adaptation during invasion.

Introduction History Influences Aboveground Biomass Allocation in Brazilian Peppertree (Schinus terebinthifolius)

Multiple introductions of an exotic species can facilitate invasion success by allowing for a wider range of expressed trait values in the adventive range. Schinus terebinthifolius (Brazilian peppertree) is an invasive shrub that was introduced into Florida in two separate introductions and has subsequently hybridized, resulting in three distinct lineages (eastern, western, and hybrid). To determine whether allocation of aboveground biomass differed by introduction history, we destructively sampled 257 stems from each of six populations with differing introduction histories. The proportion of aboveground biomass allocated to fruit, wood, and leaves differed among the three populations. To determine whether the relationship between stem size and several dependent variables that measure plant performance (total dry weight, wood dry weight, number of fruits, fruit dry weight, leaf dry weight, and number of leaves) differed quantitatively by introduction history, we performed analyses of covariance. Slopes of these relationships (dependent variable vs. stem size) varied by lineage. Hybrid populations had the steepest slopes for one set of dependent variables (total dry weight, wood dry weight, and leaf dry weight), while western populations had the steepest slopes for a different set of dependent variables (number of fruits, fruit dry weight, and number of leaves). The parameterized regression equations for each dependent variable and lineage were used to nondestructively estimate different kinds of production by individuals that are part of long-term longitudinal studies to understand the demographic consequences of these different biomass allocation strategies for the performance of S. terebinthifolius individuals across the invaded range in Florida.

Ascidians as models for studying invasion success

During the past three decades, coastal marine waters have become among the most invaded habitats globally. Ascidians are among the most notorious invaders in these ecosystems. Owing to their rapid spread, frequent population outbreaks, and associated negative ecological and economic impacts, invasive ascidians have become a global problem. Thus, the study of ascidian invasions has become a prominent area of invasion biology. Here, we review current knowledge and conclude that ascidians are good models for studying invasion success in the marine realm. Firstly, we summarize the reconstruction of invasion pathways or colonization histories and associated negative impacts of invasive ascidians, and address the urgent need to clarify ambiguous taxonomy of ascidians. Secondly, we discuss factors that underlie or facilitate invasion success of ascidians, including vectors of introduction and spread, environmental changes, biological traits, and possible genetic issues. Finally, we summarize current science-based policies and management solutions that are in place to prevent and control spread of invasive ascidians. We conclude by highlighting key research questions that remain to be answered, and propose future research to investigate mechanisms of invasion success in the marine realm using ascidians as model systems.

Chemical cues released by heterospecific competitors: behavioural responses of native and alien invasive aquatic gastropods

Chemical cues released by organisms have the potential to impact the behaviour of species, particularly in aquatic environments. These interactions affect the dynamics of resource utilisation at a small scale. Chemical cues may also be significantly involved in heterospecific competition and in reducing biotic resistance thus facilitating invasion success. This occurs if one species causes another to vacate optimal niches. This study aimed to assess (1) the behavioural responses of native gastropods to chemical cues released by other native gastropod species with which they did not co-occur, and (2) whether native gastropods from different localities exhibit similar behavioural responses to chemical cues released by the alien invasive Tarebia granifera, irrespective of their prior exposure to the alien. Behavioural responses were measured using an experimental approach both in situ and in the laboratory. Principal Components Analyses were used to visualize the variation in the behavioural responses of gastropods which were measured in terms of displacement, number of steps and turning angles. Gastropod behaviour was similar for populations from different sites, irrespective of their prior exposure to T. granifera. The behaviour of native gastropods was only different when exposed to chemical cues naturally released by T. granifera. Additionally, native gastropods did not behave differently when exposed to the chemical cues of an alien physid, Aplexa marmorata. This suggests that a change in behaviour when exposed to chemical cues of T. granifera may not be the result of having different evolutionary histories, but rather that T. granifera has evolved a unique metabolite which acts a deterrent to potential competitive snails. The role of chemical cues in mediating interactions between native and invasive species needs further research, as they may be important factors influencing biotic resistance and facilitating invasion success.

The role thermal physiology plays in species invasion.

The characterization of physiological phenotypes that may play a part in the establishment of non-native species can broaden our understanding about the ecology of species invasion. Here, an assessment was carried out by comparing the responses of invasive and native species to thermal stress. The goal was to identify physiological patterns that facilitate invasion success and to investigate whether these traits are widespread among invasive ectotherms. Four hypotheses were generated and tested using a review of the literature to determine whether they could be supported across taxonomically diverse invasive organisms. The four hypotheses are as follows: (i) broad geographical temperature tolerances (thermal width) confer a higher upper thermal tolerance threshold for invasive rather than native species; (ii) the upper thermal extreme experienced in nature is more highly correlated with upper thermal tolerance threshold for invasive vs. native animals; (iii) protein chaperone expression-a cellular mechanism that underlies an organism's thermal tolerance threshold-is greater in invasive organisms than in native ones; and (iv) acclimation to higher temperatures can promote a greater range of thermal tolerance for invasive compared with native species. Each hypothesis was supported by a meta-analysis of the invasive/thermal physiology literature, providing further evidence that physiology plays a substantial role in the establishment of invasive ectotherms.