Variation in the impact of non‐native seaweeds along gradients of habitat degradation: a meta‐analysis and an experimental test

Biological invasions are acknowledged among the main drivers of global changes in biodiversity. Despite compelling evidence of species interactions being strongly regulated by environmental conditions, there is a dearth of studies investigating how the effects of non‐native species vary among areas exposed to different anthropogenic pressures. Focusing on marine macroalgae, we performed a meta‐analysis to test whether and how the direction and magnitude of their effects on resident communities and species varies in relation to cumulative anthropogenic impact levels. The relationship between human impact levels and non‐native species impact intensity emerged only for a reduced subset of the response variables examined. Yet, there was a trend for the effects of non‐native species on community biomass and abundance and on species abundance to become less negative at heavily impacted sites. By contrast, the magnitude of negative effects of seaweed on community evenness tended to increase with human impact levels. The hypothesis of decreasing severity of invader’ impacts along a gradient of habitat degradation was also tested experimentally at a regional scale by comparing the effects of the removal of non‐native alga, Caulerpa cylindracea, on resident assemblages among rocky reefs exposed to different anthropogenic pressures. Assemblages at urban and pristine site did not differ when invaded, but did so when C. cylindracea was removed. Our results suggest that, despite the generally weak relationship between human impacts levels and non‐native species impacts, more negative impacts can be expected in less stressful environments (i.e. less degraded or pristine sites), where competitive interactions are presumably the driving force structuring resident communities. Implementing strategies for controlling the establishment of non‐native seaweeds should be, thus, considered a priority for preserving biodiversity in relatively pristine areas. On the other hand, control of invaders at degraded sites could be warranted to lessen their role as propagule sources. Synthesis Local anthropogenic stressors that severely alter biotic and abiotic conditions may underpin context‐dependency in the impacts of biological invasions. We used a meta‐analysis and an experimental test to examine the relationship between cumulative human impacts and ecological impact of non‐native seaweeds on resident assemblages. Our results suggest that more negative impacts of non‐native seaweeds on the abundance and biomass of resident assemblages can be expected in less degraded or pristine sites. Possibly, stronger impacts prevail at pristine sites, where assemblages are mainly structured by biotic interactions. Hence, management efforts should be mostly directed to prevent the establishment and spread of non‐native seaweeds in pristine areas. On the other hand, weak, but positive effects of seaweeds at the most degraded sites add to the ongoing debate on the role of non‐native species in rehabilitation plans.     

Antarctica: The final frontier for marine biological invasions

Antarctica is experiencing significant ecological and environmental change, which may facilitate the establishment of non‐native marine species. Non‐native marine species will interact with other anthropogenic stressors affecting Antarctic ecosystems, such as climate change (warming, ocean acidification) and pollution, with irreversible ramifications for biodiversity and ecosystem services. We review current knowledge of non‐native marine species in the Antarctic region, the physical and physiological factors that resist establishment of non‐native marine species, changes to resistance under climate change, the role of legislation in limiting marine introductions, and the effect of increasing human activity on vectors and pathways of introduction. Evidence of non‐native marine species is limited: just four marine non‐native and one cryptogenic species that were likely introduced anthropogenically have been reported freely living in Antarctic or sub‐Antarctic waters, but no established populations have been reported; an additional six species have been observed in pathways to Antarctica that are potentially at risk of becoming invasive. We present estimates of the intensity of ship activity across fishing, tourism and research sectors: there may be approximately 180 vessels and 500+ voyages in Antarctic waters annually. However, these estimates are necessarily speculative because relevant data are scarce. To facilitate well‐informed policy and management, we make recommendations for future research into the likelihood of marine biological invasions in the Antarctic region.     

A framework for understanding human‐driven vegetation change

Despite a major research focus on human‐mediated reshuffling of plant communities, no coherent framework unites the numerous types of changes in abundances and distributions of native and non‐native species that are driven by human activities. Human driven vegetation change can occur through: non‐native species introductions; population outbreaks or collapses; range expansions or contractions; and range shifts of both native and non‐native species. Boundaries among these different types of floristic changes are not always distinct because of an overlap in the ecological, climatic, and anthropogenic processes that underpin them. We propose a new framework that connects various human‐mediated causes of vegetation change, highlights the spatial scales at which drivers act and the temporal scale at which plant assemblages respond, and provides critical insights for identifying and appropriately managing these changes.     

Modelling marine community responses to climate‐driven species redistribution to guide monitoring and adaptive ecosystem‐based management

As a consequence of global climate‐driven changes, marine ecosystems are experiencing polewards redistributions of species – or range shifts – across taxa and throughout latitudes worldwide. Research on these range shifts largely focuses on understanding and predicting changes in the distribution of individual species. The ecological effects of marine range shifts on ecosystem structure and functioning, as well as human coastal communities, can be large, yet remain difficult to anticipate and manage. Here, we use qualitative modelling of system feedback to understand the cumulative impacts of multiple species shifts in south‐eastern Australia, a global hotspot for ocean warming. We identify range‐shifting species that can induce trophic cascades and affect ecosystem dynamics and productivity, and evaluate the potential effectiveness of alternative management interventions to mitigate these impacts. Our results suggest that the negative ecological impacts of multiple simultaneous range shifts generally add up. Thus, implementing whole‐of‐ecosystem management strategies and regular monitoring of range‐shifting species of ecological concern are necessary to effectively intervene against undesirable consequences of marine range shifts at the regional scale. Our study illustrates how modelling system feedback with only limited qualitative information about ecosystem structure and range‐shifting species can predict ecological consequences of multiple co‐occurring range shifts, guide ecosystem‐based adaptation to climate change and help prioritise future research and monitoring.     

Current knowledge on non‐native freshwater fish introductions

This review provides a contemporary account of knowledge on aspects of introductions of non‐native fish species and includes issues associated with introduction pathways, ecological and economic impacts, risk assessments, management options and impact of climate change. It offers guidance to reconcile the increasing demands of certain stakeholders to diversify their activities using non‐native fishes with the long‐term sustainability of native aquatic biodiversity. The rate at which non‐native freshwater fishes have been introduced worldwide has doubled in the space of 30 years, with the principal motives being aquaculture (39%) and improvement of wild stocks (17%). Economic activity is the principal driver of human‐mediated non‐native fish introductions, including the globalization of fish culture, whereby the production of the African cichlid tilapia is seven times higher in Asia than in most areas of Africa, and Chile is responsible for c. 30% of the world's farmed salmon, all based on introduced species. Consequently, these economic benefits need balancing against the detrimental environmental, social and economic effects of introduced non‐native fishes. There are several major ecological effects associated with non‐native fish introductions, including predation, habitat degradation, increased competition for resources, hybridization and disease transmission. Consideration of these aspects in isolation, however, is rarely sufficient to adequately characterize the overall ecological effect of an introduced species. Regarding the management of introduced non‐native fish, pre‐introduction screening tools, such as the fish invasiveness scoring kit (FISK), can be used to ensure that species are not introduced, which may develop invasive populations. Following the introduction of non‐native fish that do develop invasive populations, management responses are typified by either a remediation or a mitigation response, although these are often difficult and expensive to implement, and may have limited effectiveness.     

Non‐native plants have greater impacts because of differing per‐capita effects and nonlinear abundance–impact curves

Invasive, non‐native species can have tremendous impacts on biotic communities, where they reduce the abundance and diversity of local species. However, it remains unclear whether impacts of non‐native species arise from their high abundance or whether each non‐native individual has a disproportionate impact – that is, a higher per‐capita effect – on co‐occurring species compared to impacts by native species. Using a long‐term study of wetlands, we asked how temporal variation in dominant native and non‐native plants impacted the abundance and richness of other plants in the recipient community. Non‐native plants reached higher abundances than natives and had greater per‐capita effects. The abundance–impact relationship between plant abundance and richness was nonlinear. Compared with increasing native abundance, increasing non‐native abundance was associated with steeper declines in richness because of greater per‐capita effects and nonlinearities in the abundance–impact relationship. Our study supports eco‐evolutionary novelty of non‐natives as a driver of their outsized impacts on communities.     

Non‐native species modify the isotopic structure of freshwater fish communities across the globe

Multiple anthropogenic pressures including the widespread introductions of non‐native species threaten biodiversity and ecosystem functioning notably by modifying the trophic structure of communities. Here, we provided a global evaluation of the impacts of non‐native species on the isotopic structure (δ¹³C and δ¹⁵N) of freshwater fish communities. We gathered the stable isotope values (n = 4030) of fish species in 496 fish communities in lentic (lakes, backwaters, reservoirs) and lotic (running waters such as streams, rivers) ecosystems throughout the world and quantified the isotopic structure of communities. Overall, we found that communities containing non‐native species had a different isotopic structure than communities without non‐native species. However, these differences varied between ecosystem types and the trophic positions of non‐native species. In lotic ecosystems, communities containing non‐native species had a larger total isotopic niche than communities without non‐native species. This was primarily driven by the addition of non‐native predators at the top of the food chain that increased δ¹⁵N range without modifying the isotopic niche size of native species. In lentic ecosystems, non‐native primary consumers increased δ¹⁵N range and this was likely driven by an increase of resource availability for species at higher trophic levels, increasing food chain length. The introduction of non‐native secondary consumers at the centre of the isotopic niche of recipient communities decreased the core isotopic niche size, the δ¹³C range of recipient communities and the total isotopic niche of coexisting native species. These results suggested a modified contribution of the basal resources consumed (e.g. multi‐chain omnivory) and an increase level of competition with native species. Our results notably imply that, by affecting the isotopic structure of freshwater fish communities at a global scale, non‐native species represent an important source of perturbations that should be accounted for when investigating macro‐ecological patterns of community structure and biotic interactions.     

Co‐varying impacts of land use and non‐native brown trout on fish communities in small streams

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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.     

Global Salmonidae introductions reveal stronger ecological effects of changing intraspecific compared to interspecific diversity

The introduction of organisms within the native range of wild conspecifics is a widespread phenomenon and locally modifies patterns in intraspecific diversity. However, our knowledge of the resulting ecological effects, as opposed to those caused by invasion‐induced changes in interspecific diversity, is still limited. Here, we investigated the ecological effects of native and non‐native invaders across levels of biological organisations and recipient organisms using the global and long history introductions of salmonids. Our meta‐analysis demonstrated that the global effects of native species introductions exceeded those induced by non‐native invaders. The impacts of native invaders were primarily manifested at the individual level on wild conspecifics, but remained largely unexplored on other native organisms and at the community and ecosystem levels. Overlooked and poorly appreciated, quantifying the impacts of native invaders has important implications because human‐assisted introductions of domesticated organisms are ubiquitous and likely to proliferate in the future.     

Multi‐trophic impacts of an invasive aquatic plant

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Dynamic species distribution models reveal spatiotemporal habitat shifts in native range‐expanding versus non‐native invasive birds in an urban area

Urbanisation as a major driver of changes leads to the extinction of some species while others increase in abundance, especially non‐native species. Spatiotemporal distribution patterns of these successful species are likely to be shaped by their response and tolerance to urban features. This study assesses the anthropo‐ecological requirements of two co‐occurring bird species, the native range‐shifting jackdaw Corvus monedula and the non‐native invasive ring‐necked parakeet Psittacula krameri. We built yearly models over an eight‐year period using an ensemble modelling approach assessing response differences through time and between species. Predictors describing human‐made structures, socio‐ecological proxies and resources availability were extracted from temporally coincident databases. Dispersal and habitat constraints were implemented in final models to provide more realistic forecasts of species future distributions. Ensemble models evaluated with a random partition of the training dataset showed a higher accuracy than those evaluated with an independent dataset from another time period. Our results highlight temporal variations in the relative importance of predictors for both studied species. Single‐season occurrence data may thus be insufficient to characterize species ecological requirements. The ring‐necked parakeet and the jackdaw showed different responses to urban features. Jackdaws preferred the more urbanized part of the city while the distribution of parakeets was strongly positively associated with the density of exotic ornamental trees. We concluded that ring‐necked parakeet range expansion is likely to be driven by its effective ability to exploit urban resources which native species do not or under exploit, suggesting an open window of foraging opportunities. However, the jackdaw may be misled by a high cavity availability and a large amount of low‐quality anthropogenic food in the urban core. We suggest that dynamic SDMs are a critical tool not only to forecast the future expansion of invasive species but also for a better understanding of processes driving urban biodiversity persistence.     

Does a long‐term shift in Wood Stork diet foreshadow adaptability to human‐induced rapid environmental change?

To preserve biodiversity, identifying at‐risk populations and developing conservation plans to mitigate the effects of human‐induced rapid environmental change (HIREC) are essential. Changes in diet, especially for food‐limited species, can aid in detecting populations being impacted by HIREC, and characterizing the quality, abundance, and temporal and spatial consistency of newly consumed food items may provide insight concerning the likelihood of a species persisting in a changing environment. We used Wood Storks (Mycteria americana) nesting in the Florida Everglades as a model system to study the possible effects of HIREC on a food‐limited population. We compared the diets of Wood Storks in 2013 and 2014 with those reported during the 1970s before major anthropogenic activities affected the Everglades system and prey availability. Wood Storks in our study consumed more large‐bodied sunfish species (Lepomis spp.), fewer native marsh fishes, and more non‐native fish species than during the 1970s. Large sunfish and non‐native fish are relatively rare in the drying pools of Everglades marshes where storks traditionally forage, suggesting that Wood Storks may be using novel foraging habitats such as created wetlands (i.e., canals and stormwater ponds). Although created wetlands have long hydroperiods conducive to maintaining large‐bodied fishes and could provide alternative foraging habitat when prey availability is reduced in natural marshes, additional studies are needed to determine the extent to which these wetlands are used by Wood Storks and, importantly, the quality of prey items potentially available to foraging Wood Storks in created wetlands.     

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.     

Phenotypes of Two Generations of Sporobolus airoides Seedlings Derived from Acroptilon repens‐invaded and Non‐invaded Grass Populations

Although the ecological impacts of invasive species are well known, the evolutionary impacts on recipient native grass communities are not. We suggest that remnant native plants may provide desirable seed sources for restoration and native plant production. Native populations exposed to the selective pressures associated with exotic invasion may retain traits that increase their ability to coexist with invasive species. Two generations of Sporobolus airoides Torr. (Alkali sacaton) plants derived from lineages collected from within long‐term invaded areas of Acroptilon repens (L.) DC (Russian knapweed) and from adjacent non‐invaded areas were propagated in a greenhouse to evaluate generational changes in phenotypic traits from the production environment. Given the difference in invasion history of the two populations, we hypothesized that invaded and non‐invaded subpopulations would differ phenotypically. Phenotypic measurements revealed that invaded subpopulations had greater vegetative growth, whereas non‐invaded subpopulations had increased sexual reproduction. Phenotypic expression changed from the first to the second generation, predominantly in the invaded subpopulation. Generational phenotypic shifts are disadvantageous for native seed production which requires a standard product to sell commercially. However, phenotypic variation may improve field seed survival. This research demonstrates the potential value of targeting post‐invasion remnant grass populations for restoration.     

Time and environment explain the current richness distribution of non‐marine turtles worldwide

Ecological, historical, and evolutionary hypotheses are important to explain geographical diversity gradients in many clades, but few studies have combined them into a single analysis allowing a comparison of their relative importance. This study aimed to evaluate the relative importance of ecological, historical, and evolutionary hypotheses in explaining the current global distribution of non‐marine turtles, a group whose distribution patterns are still poorly explored. We used data from distribution range maps of 336 species of non‐marine turtles, environmental layers, and phylogeny to obtain richness estimates of these animals in 2° × 2° cells and predictors related to ecological, evolutionary and historical hypotheses driving richness patterns. Then we used a path analysis to evaluate direct and indirect effects of the predictors on turtle richness. Ancestral area reconstruction was also performed in order to evaluate the influence of time‐for‐speciation in the current diversity of the group. We found that environmental variables had the highest direct effects on non‐marine turtle richness, whereas diversification rates and area available in the last 55 million yr minimally influenced turtle distributions. We found evidence for the time‐for‐speciation effect, since regions colonized early were generally richer than recently colonized regions. In addition, regions with a high number of colonization events had a higher number of turtle species. Our results suggested that ecological processes may influence non‐marine turtle richness independent of diversification rates, but they are probably related to dispersal abilities. However, colonization time was also an important component that must be taken into account. Finally, our study provided additional support for the importance of ecological (climate and productivity) and historical (time‐for‐speciation and dispersal) processes in shaping current biodiversity patterns.     

Altered ignition catchments threaten a hyperdiverse fire‐dependent ecosystem

Human activities affect fire in many ways, often unintentionally or with considerable time‐lags before they manifest themselves. Anticipating these changes is critical, so that insidious impacts on ecosystems, their biodiversity and associated goods and services can be avoided, mitigated or managed. Here we explore the impact of anthropogenic land cover change on fire and biodiversity in adjacent ecosystems on the hyperdiverse Cape Peninsula, South Africa. We develop a conceptual framework based on the notion of an ignition catchment, or the spatial extent and temporal range where an ignition is likely to result in a site burning. We apply this concept using fire models to estimate spatial changes in burn probability between historical and current land cover. This change layer was used to predict the observed record of fires and forest encroachment into fire‐dependent Fynbos ecosystems in Table Mountain National Park. Urban expansion has created anthropogenic fire shadows that are modifying fire return intervals, facilitating a state shift to low‐diversity, non‐flammable forest at the expense of hyperdiverse, flammable Fynbos ecosystems. Despite occurring in a conservation area, these ecosystems are undergoing a hidden collapse and desperately require management intervention. Anthropogenic fire shadows can be caused by many human activities and are likely to be a universal phenomenon, not only contributing to the observed global decline in fire activity but also causing extreme fires in ecosystems where there is no shift to a less flammable state and flammable fuels accumulate. The ignition catchment framework is highly flexible and allows detection or prediction of changes in the fire regime, the threat this poses for ecosystems or fire risk and areas where management interventions and/or monitoring are required. Identifying anthropogenic impacts on ignition catchments is key for both understanding global impacts of humans on fire and guiding management of human‐altered landscapes for desirable outcomes.     

The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta‐analysis)

Globally, biological invasions can have strong impacts on biodiversity as well as ecosystem functioning. While less conspicuous than introduced aboveground organisms, introduced belowground organisms may have similarly strong effects. Here, we synthesize for the first time the impacts of introduced earthworms on plant diversity and community composition in North American forests. We conducted a meta‐analysis using a total of 645 observations to quantify mean effect sizes of associations between introduced earthworm communities and plant diversity, cover of plant functional groups, and cover of native and non‐native plants. We found that plant diversity significantly declined with increasing richness of introduced earthworm ecological groups. While plant species richness or evenness did not change with earthworm invasion, our results indicate clear changes in plant community composition: cover of graminoids and non‐native plant species significantly increased, and cover of native plant species (of all functional groups) tended to decrease, with increasing earthworm biomass. Overall, these findings support the hypothesis that introduced earthworms facilitate particular plant species adapted to the abiotic conditions of earthworm‐invaded forests. Further, our study provides evidence that introduced earthworms are associated with declines in plant diversity in North American forests. Changing plant functional composition in these forests may have long‐lasting effects on ecosystem functioning.     

Severity of impacts of an introduced species corresponds with regional eco‐evolutionary experience

Invasive plant impacts vary widely across introduced ranges. We tested the hypothesis that differences in the eco‐evolutionary experience of native communities with the invader correspond with the impacts of invasive species on native vegetation, with impacts increasing with ecological novelty. We compared plant species richness and composition beneath Pinus contorta to that in adjacent vegetation and other P. contorta stands across a network of sites in its native (Canada and USA) and non‐native (Argentina, Chile, Finland, New Zealand, Scotland, Sweden) ranges. At sites in North America and Europe, within the natural distribution of the genus Pinus, P. contorta was not associated with decreases in diversity. In the Southern Hemisphere, where there are no native Pinaceae, plant communities beneath P. contorta were less diverse than in other regions and compared to uninvaded native vegetation. Effects on native vegetation were particularly pronounced where P. contorta was a more novel life form and exhibited higher growth rates. Our results support the hypothesis that the eco‐evolutionary experience of the native vegetation, and thus the novelty of the invader, determines the magnitude of invader impacts on native communities. Understanding the eco‐evolutionary context of invasions will help to better understand and predict where invasion impacts will be greatest and to prioritize invasive species management.