Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.     

Differential temporal beta‐diversity patterns of native and non‐native arthropod species in a fragmented native forest landscape

An important factor that hinders the management of non‐native species is a general lack of information regarding the biogeography of non‐natives, and, in particular, their rates of turnover. Here, we address this research gap by analysing differences in temporal beta‐diversity (using both pairwise and multiple‐time dissimilarity metrics) between native and non‐native species, using a novel time‐series dataset of arthropods sampled in native forest fragments in the Azores. We use a null model approach to determine whether temporal beta‐diversity was due to deterministic processes or stochastic colonisation and extinction events, and linear modelling selection to assess the factors driving variation in temporal beta‐diversity between plots. In accordance with our predictions, we found that the temporal beta‐diversity was much greater for non‐native species than for native species, and the null model analyses indicated that the turnover of non‐native species was due to stochastic events. No predictor variables were found to explain the turnover of native or non‐native species. We attribute the greater turnover of non‐native species to source‐sink processes and the close proximity of anthropogenic habitats to the fragmented native forest plots sampled in our study. Thus, our findings point to ways in which the study of turnover can be adapted for future applications in habitat island systems. The implications of this for biodiversity conservation and management are significant. The high rate of stochastic turnover of non‐native species indicates that attempts to simply reduce the populations of non‐native species in situ within native habitats may not be successful. A more efficient management strategy would be to interrupt source‐sink dynamics by improving the harsh boundaries between native and adjacent anthropogenic habitats.     

Periphyton density is similar on native and non‐native plant species

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Predicting the outcomes of a tri‐trophic interaction between an indigenous parasitoid and an exotic herbivorous pest and its host plants

Understanding the novel ecological interactions that result from biological invasions is a critical issue in modern ecology and evolution as well as pest management. Introduced herbivorous insects may interact with native plants and indigenous natural enemies, creating novel tri‐trophic interactions. To help predict the potential outcomes of novel interactions, we investigated the behavioural and physiological responses of an indigenous generalist parasitoid (Habrobracon gelechiae) to an introduced generalist herbivore (the light brown apple moth, Epiphyas postvittana) and its new host plants in California. We first examined the parasitoid's host location and acceptance on a range of nine common host plants of the moth representing distinctly different geographic origins and morphologies (to examine the effect of a known toxic plant on the parasitoid's performance, an additional toxic plant species was also tested that the moth consumes in the laboratory but does not naturally attack). The parasitoid was able to locate the host larvae on all plants equally well, although clutch size was affected by host plant. We then determined fitness of the moth and the parasitoid on four representative plants. The moth larvae suffered higher mortality and a slower developmental rate on the known toxic plant than on the other three plants, but the parasitoid's fitness correlates did not differ between the host food plants. These results show a high level of plasticity in the indigenous generalist parasitoid in its ability to exploit the exotic host on a wide range of host plants, generating an invasion‐driven novel tri‐trophic interaction.     

A novel environmental DNA approach to quantify the cryptic invasion of non‐native genotypes

The invasion of non‐native species that are closely related to native species can lead to competitive elimination of the native species and/or genomic extinction through hybridization. Such invasions often become serious before they are detected, posing unprecedented threats to biodiversity. A Japanese native strain of common carp (Cyprinus carpio) has become endangered owing to the invasion of non‐native strains introduced from the Eurasian continent. Here, we propose a rapid environmental DNA‐based approach to quantitatively monitor the invasion of non‐native genotypes. Using this system, we developed a method to quantify the relative proportion of native and non‐native DNA based on a single‐nucleotide polymorphism using cycling probe technology in real‐time PCR. The efficiency of this method was confirmed in aquarium experiments, where the quantified proportion of native and non‐native DNA in the water was well correlated to the biomass ratio of native and non‐native genotypes. This method provided quantitative estimates for the proportion of native and non‐native DNA in natural rivers and reservoirs, which allowed us to estimate the degree of invasion of non‐native genotypes without catching and analysing individual fish. Our approach would dramatically facilitate the process of quantitatively monitoring the invasion of non‐native conspecifics in aquatic ecosystems, thus revealing a promising method for risk assessment and management in biodiversity conservation.     

A modified DNA barcode approach to define trophic interactions between native and exotic pentatomids and their parasitoids

The establishment of invasive Halyomorpha halys (Stål) outside of its native range may impact native species assemblages, including other pentatomids and their scelionid parasitoids. This has generated interest in defining species diversity and host‐parasitoid associations in this system to better understand the impact of invasive alien species on trophic interactions in invaded regions. Information on scelionid–pentatomid associations in natural habitats is lacking, and species‐level identification of these associations can be tenuous using rearing and dissection techniques. Naturally occurring pentatomid eggs were collected in areas where H. halys has established in Canada and were analysed using a modified DNA barcoding approach to define species‐level trophic interactions. Identification was possible for >90% of egg masses. Eleven pentatomid and five scelionid species were identified, and trophic links were established. Approximately 70% of egg masses were parasitized; parasitism and parasitoid species composition were described for each species. Telenomus podisi Ashmead was the dominant parasitoid and was detected in all host species. Trissolcus euschisti Ashmead was detected in several host species, but was significantly more prevalent in Chinavia hilaris (Say) and Brochymena quadripustulata (Fabricius). Trissolcus brochymenae Ashmead and Tr. thyantae Ashmead were recorded sporadically. Parasitism of H. halys was 55%, and this species was significantly less likely to be parasitized than native pentatomids. The scelionid species composition of H. halys consisted of Te. podisi, Tr. euschisti and Tr. thyantae. Although these species cannot develop in fresh H. halys eggs, we demonstrate that parasitoids attempt to exploit this host under field conditions.     

Landscape characteristics of non‐native pine plantations and invasions in Southern Chile

The spread of non‐native conifers into areas naturally dominated by other vegetation types is a growing problem in South America. This process results in a landscape transformation as the conifers suppress native vegetation leading to reduced biodiversity, lower water availability and altered nutrient dynamics. Previous research highlights the broad spatial extents of land cover change in parts of Chile. However, in Southern Chile, the extent of plantations and the landscape characteristics associated with plantations and ongoing pine invasions are poorly understood. Here, we characterised non‐native pine land cover within one Landsat scene (World Reference System 2 Path 232/Row 92; ~34 000 km²) in Southern Chile. We created training data based on historical high‐resolution imagery, derived land cover predictors from time series of Landsat observations and used a Random Forest classifier to map the distribution of non‐native pines. The overall classification accuracy was 88%, and the accuracy of the non‐native pine class exceeded 90%. Although 71% of non‐native pine patches were within 500 m of other non‐native pine patches, isolated non‐native pine patches were found to occur up to 55 km from the nearest neighbour. These distant plantations could exacerbate invasion risk by creating propagule sources for novel invasion fronts. In relation to landscape characteristics, non‐native pines were found to be more likely to occur in low slope and mid‐elevation areas. Because most of the study area is native forest, most non‐native pine patches border native forest. However, non‐native pine patches were almost three times more likely than random patches to border grass/agriculture. This suggests that grasslands and disturbed sites, which have low resistance to non‐native pine invasion, are disproportionately exposed to pine propagules. Our results indicate that non‐native pine plantations are extensive across Southern Chile, and well poised to cause future invasion.     

Presence and absence of non‐native fish species in the Wet Tropics region,Australia

Distributional records of non‐native fish species were identified in the Wet Tropics region, Far North Queensland, Australia, through a compilation of published records and expert knowledge. A total of 1106 records were identified comprising 346 presence and four uncertain records for at least 13 species, and 756 absence records. All current presence records consist of six species from the families Cichlidae and Poeciliidae with established self‐sustaining populations in the region, probably affecting the highly diverse native fish fauna.     

Thermal adaptation affects interactions between a range‐expanding and a native odonate species

1. Increasing temperature and invading species may interact in their effects on communities. In this study, we investigated how rising temperatures alter larval interactions between a naturally range‐expanding dragonfly, Crocothemis erythraea, and a native northern European species, Leucorrhinia dubia. Initial studies revealed that C. erythraea grow up to 3.5 times faster than L. dubia at temperatures above 16 °C. As a result, we hypothesised that divergent temperature responses would lead to rapid size differences between coexisting larvae and, consequently, to asymmetric intraguild predation at higher ambient temperatures. 2. Mortality and growth rates were measured in interaction treatments (with both species present) and non‐interaction controls (one species present) at four different temperature regimes: at an ambient temperature representative of central Germany, where both species overlap in distribution, and at temperatures increased by 2, 4 and 6 °C. 3. The mortality of C. erythraea did not differ between treatment and control. In contrast, mortality of L. dubia remained similar over all temperatures in the controls, but increased with temperature in the presence of the other species and was significantly higher there than in the controls. We concluded that L. dubia suffered asymmetric intraguild predation, particularly at increased temperature. Reduced growth rate of L. dubia in the interaction treatment at higher temperatures also suggested asymmetric competition for prey in the first phase of the experiment. 4. The results imply that the range expansion of C. erythraea may cause reduction in population size of syntopic L. dubia when temperature rises by more than 2 °C. The consequences for future range patterns, as well as other factors that may influence the interaction in nature, are discussed.     

Selection on tropane alkaloids in native and non‐native populations of Datura stramonium

Theories of plant invasion based on enemy release in a new range assume that selection exerted by specialist herbivores on defence traits should be reduced, absent, or even selected against in the new environment. Here, we measured phenotypic selection on atropine and scopolamine concentration of Datura stramonium in eight native (Mexico) and 14 non‐native (Spain) populations. Native populations produced between 20 and 40 times more alkaloid than non‐native populations (atropine: 2.0171 vs. 0.0458 mg/g; scopolamine: 1.004 vs. 0.0488 mg/g, respectively). Selection on alkaloids was negative for atropine and positive for scopolamine concentration in both ranges. However, the effect sizes of selection gradients were only significant in the native range. Our results support the assumption that the reduction of plant defence in the absence of the plant's natural enemies in invasive ranges is driven by natural selection.     

Morphological variability of black bullhead Ameiurus melas in four non‐native European populations

External morphology in black bullhead Ameiurus melas, a fish species considered to have high invasive potential, was studied in its four non‐native European populations (British, French, Italian and Slovak). The aim of this study was to examine this species' variability in external morphology, including ontogenetic context, and to evaluate its invasive potential. Specimens from all non‐native populations reached smaller body size compared to individuals from native populations. Juvenile A. melas were found to have a relatively uniform body shape regardless of the population's origin, whereas adults developed different phenotypes depending upon location. Specimens from the U.K., Slovak and French populations appeared to be rather similar to each other, whereas the Italian population showed the most distant phenotype. This probably results from the different thermal regime in the Italian habitat. Ameiurus melas from non‐native European populations examined in this study showed some potential to alter the body shape both within and between populations. The phenotypic plasticity of A. melas, however, was not found to be as significant as in other invasive fish species. The results suggest that morphological variability itself is not necessarily essential for invasive success. The invasiveness of A. melas is therefore probably favoured by variations in its life‐history traits and reproduction variables, together with some behavioural traits (e.g. voracious feeding and parental care) rather than by phenotypic plasticity expressed in external morphology.     

Insect diversity and trophic structure differ on native and non‐indigenous congeneric rushes in coastal salt marshes

Displacement of native plant species by non‐indigenous congeners may affect associated faunal assemblages. In endangered salt marshes of south‐east Australia, the non‐indigenous rush Juncus acutus is currently displacing the native rush Juncus kraussii, which is a dominant habitat‐forming species along the upper border of coastal salt marshes. We sampled insect assemblages on multiple plants of these congeneric rushes in coastal salt marshes in Sydney, New South Wales, Australia, and compared the abundance, richness, diversity, composition and trophic structure between: (i) J. acutus and J. kraussii at invaded locations; and (ii) J. kraussii at locations either invaded or not invaded by J. acutus. Although J. acutus supported a diverse suite of insects, species richness and diversity were significantly greater on the native J. kraussii. Moreover, insect assemblages associated with J. kraussii at sites invaded by J. acutus were significantly different from, and more variable than, those on J. kraussii at non‐invaded sites. The trophic structure of the insect assemblages was also different, including the abundance and richness of predators and herbivores, suggesting that J. acutus may be altering consumer interactions, and may be spreading in part because of a reduction in herbivory. This strongly suggests that J. acutus is not playing a functionally similar role to J. kraussii with respect to the plant‐associated insect species assemblages. Consequently, at sites where this non‐indigenous species successfully displaces the native congener, this may have important ecological consequences for community composition and functioning of these endangered coastal salt marshes.     

Testing multiple pathways for impacts of the non‐native Black‐headed Weaver Ploceus melanocephalus on native birds in Iberia in the early phase of invasion

Not all non‐native species have strong negative impacts on native species. It is desirable to assess whether a non‐native species will have a negative impact at an early stage in the invasion process, when management options such as eradication are still available. Although it may be difficult to detect early impacts of non‐native species, it is necessary to ensure that management decisions can be based on case‐specific scientific evidence. We assess the impacts of a non‐native bird, the Black‐headed Weaver Ploceus melanocephalus, at an early stage in its invasion of the Iberian Peninsula. To do this we identify potential pathways by which competition for shared resources by Black‐headed Weavers could lead to population declines in two ecologically similar native species, and generate hypotheses to test for evidence of competition along these pathways. Black‐headed Weavers could potentially impact native species by displacing them from nesting habitat, or by reducing habitat quality. We found no evidence for either potential competition pathway, suggesting that Black‐headed Weavers do not currently compete with the two native species. However, it is possible that mechanisms that currently allow coexistence may not operate once Black‐headed Weavers reach higher population densities or different habitats.     

Global meta‐analysis of native and nonindigenous trophic traits in aquatic ecosystems

Ecologists have recently devoted their attention to the study of species traits and their role in the establishment and spread of nonindigenous species (NIS). However, research efforts have mostly focused on studies of terrestrial taxa, with lesser attention being dedicated to aquatic species. Aquatic habitats comprise of interconnected waterways, as well as exclusive introduction vectors that allow unparalleled artificial transport of species and their propagules. Consequently, species traits that commonly facilitate biological invasions in terrestrial systems may not be as represented in aquatic environments. We provide a global meta‐analysis of studies conducted in both marine and freshwater habitats. We selected studies that conducted experiments with native and NIS under common environmental conditions to allow detailed comparisons among species traits. In addition, we explored whether different factors such as species relatedness, functional feeding groups, latitude, climate, and experimental conditions could be linked to predictive traits. Our results show that species with traits that enhance consumption and growth have a substantially increased probability of establishing and spreading when entering novel ecosystems. Moreover, traits associated with predatory avoidance were more prevalent in NIS and therefore favour invasive species in aquatic habitats. When we analysed NIS interacting with taxonomically distinctive native taxa, we found that consumption and growth were particularly important traits. This suggests that particular attention should be paid to newly introduced species for which there are no close relatives in the local biota. Finally, we found a bias towards studies conducted in temperate regions, and thus, more studies in other climatic regions are needed. We conclude that studies aiming at predicting future range shifts should consider trophic traits of aquatic NIS as these traits are indicative of multiple interacting mechanisms involved in promoting species invasions.     

Niche shift in four non‐native estrildid finches and implications for species distribution models

Non‐native species can have severe impacts on ecosystems. Therefore, predictions of potentially suitable areas that are at risk of the establishment of non‐native populations are desirable. In recent years, species distribution models (SDMs) have been widely applied for this purpose. However, the appropriate selection of species records, whether from the native area alone or also from the introduced range, is still a matter of debate. We combined analyses of native and non‐native realized climate niches to understand differences between models based on all locations, as well as on locations from the native range only. Our approach was applied to four estrildid finch species that have been introduced to many regions around the world. Our results showed that SDMs based on location data from native areas alone may underestimate the potential distribution of a given species. The climatic niches of species in their native ranges differed from those of their non‐native ranges. Niche comparisons resulted in low overlap values, indicating considerable niche shifts, at least in the realized niches of these species. All four species have high potential to spread over many tropical and subtropical areas. However, transferring these results to temperate areas has a high degree of uncertainty, and we urge caution when assessing the potential spread of tropical species that have been introduced to higher latitudes.     

Feeding preferences of an invasive Ponto‐Caspian goby for native and non‐native gammarid prey

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