Performance of non-native species within marine reserves

Conservation of biodiversity is a major aim of marine reserves; however the effects of reserves on non-native species, a major threat to biodiversity globally, is not widely known. Marine reserves could resist non-native species due to enhanced native diversity and biomass that heightens biotic resistance. Or non-native species could be enhanced by reserves by at least three mechanisms, including protection from harvesting, increased fishing pressure outside reserves facilitating invasions at a regional scale and increasing the exposure of reserves to more potential invaders, and increased propagule pressure from human visitation. We exhaustively searched the literature and found 13 cases that contained quantitative data on non-native species inside and outside marine reserves. In no cases did reserves resist non-native species. Of the seven cases where reserves were established prior to the arrival of the non-native species, five had no effect on the non-native species and two enhanced non-native species. Of the six cases where reserves were established in areas that had pre-existing non-native species, two had no effect on the non-native species and four enhanced the non-native species. These results suggest that while non-native species do equally well or better within marine reserves, too few data are currently available to draw broad, general conclusions regarding the effects of marine reserves on non-native species. Management plans for marine reserves rarely include guidelines for preventing or managing non-native species. If the trends we have detected here are supported by future studies, non-native species should be a priority for management of marine reserves.     

Correlated Non-native Species Richness of Birds, Mammals, Herptiles and Plants: Scale Effects of Area, Human Population and Native Plants

Several extrinsic factors (area, native species diversity, human population size and latitude) significantly influence the non-native species richness of plants, over several orders of magnitude. Using several data sets, I examine the role of these factors in non-native species richness of several animal groups: birds, mammals and herptiles (amphibians, reptiles). I also examine if non-native species richness is correlated among these groups. I find, in agreement with Sax [2001, Journal of Biogeography 28: 139–150], that latitude is inversely correlated with non-native species richness of many groups. Once latitude is accounted for, area, human population size and native plant species richness are shown to be important extrinsic factors influencing non-native animal species. Of these extrinsic factors, human population size and native plant species richness are the best predictors of non-native animal species richness. Area, human population size and native plant species richness are highly intercorrelated, along with non-native species richness of all taxa. Indeed a factor analysis shows that a single multivariate axis explains over half of the variation for all variables among the groups. One reason for this covariation is that humans tend to most densely occupy the most productive and diverse habitats where native plant species richness is very high. It is thus difficult to disentangle the effects of human population size and native species richness on non-native species richness. However, it seems likely that these two factors may combine to increase non-native species richness in a synergistic way: high native species richness reflects greater habitat variety available for non-native species, and dense human populations (that preferentially occupy areas rich in native species) increase non-native species importation and disturbance of local habitats.     

An Anthropogenic Habitat Facilitates the Establishment of Non-Native Birds by Providing Underexploited Resources

Anthropogenic modification of habitats may reduce the resources available for native species, leading to population declines and extinction. These same habitats often have the highest richness of non-native species. This pattern may be explained if recently human-modified habitats provide novel resources that are more accessible to non-native species than native species. Using non-native birds in the Iberian Peninsula as a case study, we conduct a large-scale study to investigate whether non-native species are positively associated with human modified habitats, and to investigate whether this positive association may be driven by the presence of resources that are not fully exploited by native species. We do this by comparing the functional diversity and resource use of native and non-native bird communities in a recently human-modified habitat (rice fields) and in more traditional habitats in the Iberian Peninsula. The functional diversity of native bird communities was lower in rice fields, but non-native birds were positively associated with rice fields and plugged this gap. Differences in resource use between native and non-native species allowed non-native species to exploit resources that were plentiful in rice fields, supporting the role of underexploited resources in driving the positive association of non-native birds with rice fields. Our results provide a potential mechanism explaining the positive association of non-native species with anthropogenic habitats, and further work is needed to test if this applies more generally.     

A Framework for Evaluating Heterogeneity and Landscape-Level Impacts of Non-native Aquatic Species

Non-native species are a major component of global environmental change, and aquatic systems are especially vulnerable to non-native species impacts. Much of the research on aquatic non-native species impact has occurred at the local or site level. In reality, non-native species impacts play out across multiple spatial scales on heterogeneous landscapes. How can we ‘scale up’ our understanding of site-level impacts to the broader landscape scale? To address this disconnect, we synthesize our current understanding of key components of landscape-scale non-native species impacts: geographic range, abundance, and local impacts. Most aquatic non-native species have small ranges, while a few have large ranges. However, aquatic non-native species are often far from saturated on landscapes, and occurrence records are often woefully incomplete. Aquatic non-native species are often at low abundances where they are present, reaching high abundance in a small number of locations. Finally, local-scale impact can be estimated from abundance, but this requires knowledge of the abundance–impact relationship. Considering these multiple components enables understanding of non-native species impacts at broader spatial scales. Although the landscape-level impacts of aquatic non-native species may be high, the spatial distribution of site-level impacts is uneven, and highly impacted sites may be relatively uncommon. This heterogeneity in impacts provides an opportunity to optimize and prioritize non-native species management and prevention efforts.     

Spatial and temporal dynamics of a freshwater eukaryotic plankton community revealed via 18S rRNA gene metabarcoding

The Brazilian Environmental Ministry (MMA) released a report in 2016 listing 163 non-native species into Brazilian inland waters. Reservoirs are among the freshwater habitats most frequently associated with the release of non-native species. Therefore, the aim of the present study was to evaluate the occurrence and distribution of the non-native species listed by the MMA, in large Brazilian reservoirs (≥ 30 km²). Further, we have tested the hypothesis that beta diversity of non-native species within reservoirs from the same basin is lower than those among reservoirs from distinct basins. A literature search was performed for 70 Brazilian reservoirs, resulting in the records of 91 non-native species. Reservoirs from the Paraná basin showed the highest number of occurrences, with 33 non-native species recorded only in the Itaipu reservoir. Beta diversity of non-native species showed higher variability among reservoirs from different basins than those within the same basin. Some basins were also distinguished by their composition of non-native species, as supported by the IndVal index. Non-native species were widespread along Brazilian reservoirs, and their distribution can be even higher than reported.     

Origin matters: widely distributed native and non-native species benefit from different functional traits

Recently, ecologists debated whether distinguishing native from non-native species is sensible or not. One argument is that widespread and less widespread species are functionally different, whether or not they are native. An opposing statement points out ecologically relevant differences between native and non-native species. We studied the functional traits that drive native and non-native vascular plant species frequency in Germany by explaining species grid-cell frequency using traits and their interaction with status. Native and non-native species frequency was equally driven by life span, ploidy type and self-compatibility. Non-native species frequency rose with later flowering cessation date, whereas this relationship was absent for native species. Native and non-native species differed in storage organs and in the number of environmental conditions they tolerate. We infer that environmental filters drive trait convergence of native and non-native species, whereas competition drives trait divergence. Meanwhile, introduction pathways functionally bias the frequency of non-native species.     

Diversity and Biotic Homogenization of Urban Land-Snail Faunas in Relation to Habitat Types and Macroclimate in 32 Central European Cities

The effects of non-native species invasions on community diversity and biotic homogenization have been described for various taxa in urban environments, but not for land snails. Here we relate the diversity of native and non-native land-snail urban faunas to urban habitat types and macroclimate, and analyse homogenization effects of non-native species across cities and within the main urban habitat types. Land-snail species were recorded in seven 1-ha plots in 32 cities of ten countries of Central Europe and Benelux (224 plots in total). Each plot represented one urban habitat type characterized by different management and a specific disturbance regime. For each plot, we obtained January, July and mean annual temperature and annual precipitation. Snail species were classified into either native or non-native. The effects of habitat type and macroclimate on the number of native and non-native species were analysed using generalized estimating equations; the homogenization effect of non-native species based on the Jaccard similarity index and homogenization index. We recorded 67 native and 20 non-native species. Besides being more numerous, native species also had much higher beta diversity than non-natives. There were significant differences between the studied habitat types in the numbers of native and non-native species, both of which decreased from less to heavily urbanized habitats. Macroclimate was more important for the number of non-native than native species; however in both cases the effect of climate on diversity was overridden by the effect of urban habitat type. This is the first study on urban land snails documenting that non-native land-snail species significantly contribute to homogenization among whole cities, but both the homogenization and diversification effects occur when individual habitat types are compared among cities. This indicates that the spread of non-native snail species may cause biotic homogenization, but it depends on scale and habitat type.     

Contrasting patterns of intraspecific trait variability in native and non-native plant species along an elevational gradient on Tenerife,Canary Islands

Background and AimsNon-native plant species are not restricted to lowlands, but increasingly are invading high elevations. While for both native and non-native species we expected variability of plant functional traits due to the changing environmental conditions along elevational gradients, we additionally assumed that non-native species are characterized by a more acquisitive growth strategy, as traits reflecting such a strategy have been found to correlate with invasion success. Furthermore, the typical lowland introduction of non-native species coming from multiple origins should lead to higher trait variability within populations of non-native species specifically at low elevations, and they might therefore occupy a larger total trait space.MethodsAlong an elevational gradient ranging from 55 to 1925 m a.s.l. on Tenerife, we collected leaves from eight replicate individuals in eight evenly distributed populations of five native and six non-native forb species. In each population, we measured ten eco-morphological and leaf biochemical traits and calculated trait variability within each population and the total trait space occupied by native and non-native species.Key ResultsWe found both positive (e.g. leaf dry matter content) and negative (e.g. leaf N) correlations with elevation for native species, but only few responses for non-native species. For non-native species, within-population variability of leaf dry matter content and specific leaf area decreased with elevation, but increased for native species. The total trait space occupied by all non-native species was smaller than and a subset of that of native species.ConclusionsWe found little evidence that intraspecific trait variability is associated with the success of non-native species to spread towards higher elevations. Instead, for non-native species, our results indicate that intermediate trait values that meet the requirements of various conditions are favourable across the changing environmental conditions along elevational gradients. As a consequence, this might prevent non-native species from overcoming abruptly changing environmental conditions, such as when crossing the treeline.     

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.     

Non-native Fish Occurrence and Biomass in 1943 Western Palearctic Lakes and Reservoirs and their Abiotic and Biotic Correlates

Invasion of non-native species is considered a major threat to global biodiversity. Here we present a comprehensive overview of the occurrence, richness and biomass contribution of non-native fish species in 1943 standing water bodies from 14 countries of the Western Palearctic, based on standardised fish catches by multi-mesh gillnetting. We expected strong geographical gradients to emerge in the occurrence of non-natives. We further hypothesised that the contribution by non-natives to the local fish community biomass was correlated with local richness and the trophic level of native and non-native species. Non-native fish species occurred in 304 of 1943 water bodies (16%). If the average number of occupied water bodies per country was weighted by number of water bodies per country, the grand mean occurrence of non-natives in Western Palearctic water bodies was 10%. Exotic (non-native to the Palearctic) and translocated (non-native only to parts of the Palearctic) species were found in 164 (8.4%) or 235 (12.1%) of the water bodies, respectively. The occurrence and local richness of non-native fish species increased with temperature, precipitation and lake area and were substantially higher in reservoirs than in natural lakes. High local biomass contributions of non-native species were strongly correlated with low richness of native species and high richness of non-native species, whereas the trophic level of the fish species had only a weak effect. Single non-native species rarely dominated community biomass, but high biomass contributions and thus strong community and ecosystem impacts can be expected if several non-native species accumulate in a water body.     

Non-native fishes and native species diversity in freshwater fish assemblages across the United States

The proliferation of non-native species in North American freshwater ecosystems is considered a primary threat to the integrity of native community structure. However, a general understanding of consistent and predictable impacts of non-native species on native freshwater diversity is limited, in part, because of a lack of broad-scale studies including data from numerous localities across multiple drainages. This study uses data from 751 localities collected during the United States Geological Survey (USGS) National Water Quality Assessment (NAWQA) program to examine the influence of non-native fish species on native freshwater fish assemblages across the United States. In general, no significant differences in native fish richness and diversity measures were detected between sites with only native species and sites containing non-native species. However, at sites with non-native species, the number of non-native species present was negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Non-native piscivores were negatively correlated with native species richness and Shannon diversity and positively correlated with native evenness. Native piscivores were positively correlated with native richness and diversity and negatively correlated with native evenness at sites with only native species. Our results suggest that from a superficial perspective, native species richness and diversity are not different among sites with and without non-native species. However, when patterns of native species richness and diversity are examined at sites containing non-native species, correlations between non-native and native species richness and diversity imply the expected negative effect of invasive taxa. Additionally, non-native piscivores appear to have a significant negative effect on native taxa and possibly represent a novel selective force on naive native prey.     

Long-term trajectories of non-native vegetation on islands globally

Human-mediated changes in island vegetation are, among others, largely caused by the introduction and establishment of non-native species. However, data on past changes in non-native plant species abundance that predate historical documentation and censuses are scarce. Islands are among the few places where we can track human arrival in natural systems allowing us to reveal changes in vegetation dynamics with the arrival of non-native species. We matched fossil pollen data with botanical status information (native, non-native), and quantified the timing, trajectories and magnitude of non-native plant vegetational change on 29 islands over the past 5000 years. We recorded a proportional increase in pollen of non-native plant taxa within the last 1000 years. Individual island trajectories are context-dependent and linked to island settlement histories. Our data show that non-native plant introductions have a longer and more dynamic history than is generally recognized, with critical implications for biodiversity baselines and invasion biology.     

Non-native Ants Are Smaller than Related Native Ants

I compare the sizes of non-native and native ants to evaluate how worker size may be related to the ability of a species to invade new habitats. I compare the size of 78 non-native ant species belonging to 26 genera with the size of native congeneric species; native ants are larger than non-native ants in 22 of 26 genera. Ants were sorted by genera into fighting and nonfighting groups, based on observations of interspecific interactions with other ant species. In all of the genera with monomorphic worker castes that fight during competition, the non-native species were smaller than the native species. The genera that engage in combat had a higher frequency of significantly smaller size in non-native ants. I selected Wasmannia auropunctata for further studies, to compare native and non-native populations. Specimens of W. auropunctata from non-native populations were smaller than conspecific counterparts from its native habitat. I consider hypotheses to explain why non-native ants are smaller in size than native ants, including the role of colony size in interspecific fights, changes in life history, the release from intraspecific fighting, and climate. The discovery that fighting non-natives are smaller than their closest native relatives may provide insight into the mechanisms for success of non-native species, as well as the role of worker size and colony size during interspecific competition.     

Non-native plant invasions of United States National Parks

The United States National Park Service was created to protect and make accessible to the public the nation’s most precious natural resources and cultural features for present and future generations. However, this heritage is threatened by the invasion of non-native plants, animals, and pathogens. To evaluate the scope of invasions, the USNPS has inventoried non-native plant species in the 216 parks that have significant natural resources, documenting the identity of non-native species. We investigated relationships among non-native plant species richness, the number of threatened and endangered plant species, native species richness, latitude, elevation, park area and park corridors and vectors. Parks with many threatened and endangered plants and high native plant species richness also had high non-native plant species richness. Non-native plant species richness was correlated with number of visitors and kilometers of backcountry trails and rivers. In addition, this work reveals patterns that can be further explored empirically to understand the underlying mechanisms.     

外来种对生物多样性的影响及其控制

外来种是那些借助自身力量或其它外界力量传播到其未曾分布过的地域 ,并且能进行繁殖传播的生物。外来种入侵已成为一种引人关注的现象。外来种通过竞争、捕食、牧食、改变生境和传播疾病等方式对本地生物产生威胁 ,影响本地生物多样性。外来种入侵成了生物多样性丧失的两个主要影响之一 (另一影响是生境的破坏 )。因此 ,探寻阻止或减少外来种入侵的方法很有必要。目前 ,人们运用机械法、化学法和生物控制法来控制外来种。在外来种入侵的初始阶段或外来种数量不多时 ,运用机械法较好。化学法有带来新环境污染的危险。生物控制法是应用天敌来防治或消灭有害生物。在外来种的控制上 ,生物控制法有一些成功的案例 ,但它并不是万能的。有些生物控制剂可能对非目标种产生影响 ,这要求在释放前对生物控制剂作更严格的检测。     

Positive Effects of Non-Native Grasses on the Growth of a Native Annual in a Southern California Ecosystem

Fire disturbance is considered a major factor in the promotion of non-native plant species. Non-native grasses are adapted to fire and can alter environmental conditions and reduce resource availability in native coastal sage scrub and chaparral communities of southern California. In these communities persistence of non-native grasses following fire can inhibit establishment and growth of woody species. This may allow certain native herbaceous species to colonize and persist beneath gaps in the canopy. A field manipulative experiment with control, litter, and bare ground treatments was used to examine the impact of non-native grasses on growth and establishment of a native herbaceous species, Cryptantha muricata. C. muricata seedling survival, growth, and reproduction were greatest in the control treatment where non-native grasses were present. C. muricata plants growing in the presence of non-native grasses produced more than twice the number of flowers and more than twice the reproductive biomass of plants growing in the treatments where non-native grasses were removed. Total biomass and number of fruits were also greater in the plants growing in the presence of non-native grasses. Total biomass and reproductive biomass was also greater in late germinants than early germinants growing in the presence of non-native grasses. This study suggests a potential positive effect of non-native grasses on the performance of a particular native annual in a southern California ecosystem.     

Contrasting patterns of herbivore and predator pressure on invasive and native plants

Invasive non-native plant species often harbor fewer herbivorous insects than related native plant species. However, little is known about how herbivorous insects on non-native plants are exposed to carnivorous insects, and even less is known on plants that have recently expanded their ranges within continents due to climate warming. In this study we examine the herbivore load (herbivore biomass per plant biomass), predator load (predator biomass per plant biomass) and predator pressure (predator biomass per herbivore biomass) on an inter-continental non-native and an intra-continental range-expanding plant species and two congeneric native species. All four plant species co-occur in riparian habitat in north-western Europe. Insects were collected in early, mid and late summer from three populations of all four species. Before counting and weighing the insects were classified to trophic guild as carnivores (predators), herbivores, and transients. Herbivores were further subdivided into leaf-miners, sap-feeders, chewers and gallers. Total herbivore loads were smaller on inter-continental non-native and intra-continental range-expanding plants than on the congeneric natives. However, the differences depended on time within growing season, as well as on the feeding guild of the herbivore. Although the predator load on non-native plants was not larger than on natives, both non-native plant species had greater predator pressure on the herbivores than the natives. We conclude that both these non-native plant species have better bottom-up as well as top-down control of herbivores, but that effects depend on time within growing season and (for the herbivore load) on herbivore feeding guild. Therefore, when evaluating insects on non-native plants, variation within season and differences among feeding guilds need to be taken into account.     

Impact of Non-Native Terrestrial Mammals on the Structure of the Terrestrial Mammal Food Web of Newfoundland,Canada

The island of Newfoundland is unique because it has as many non-native terrestrial mammals as native ones. The impacts of non-native species on native flora and fauna can be profound and invasive species have been identified as one of the primary drivers of species extinction. Few studies, however, have investigated the effects of a non-native species assemblage on community and ecosystem properties. We reviewed the literature to build the first terrestrial mammal food web for the island of Newfoundland and then used network analyses to investigate how the timing of introductions and trophic position of non-native species has affected the structure of the terrestrial mammal food web in Newfoundland. The first non-native mammals (house mouse and brown rat) became established in Newfoundland with human settlement in the late 15^th and early 16^th centuries. Coyotes and southern red-backed voles are the most recent mammals to establish themselves on the island in 1985 and 1998, respectively. The fraction of intermediate species increased with the addition of non-native mammals over time whereas the fraction of basal and top species declined over time. This increase in intermediate species mediated by non-native species arrivals led to an overall increase in the terrestrial mammal food web connectance and generality (i.e. mean number of prey per predator). This diverse prey base and sources of carrion may have facilitated the natural establishment of coyotes on the island. Also, there is some evidence that the introduction of non-native prey species such as the southern red-backed vole has contributed to the recovery of the threatened American marten. Long-term monitoring of the food web is required to understand and predict the impacts of the diverse novel interactions that are developing in the terrestrial mammal food web of Newfoundland.     

Patterns of invasion of an urban remnant of a species-rich grassland in southeastern Australia by non-native plant species

Abstract. The invasion by non-native plant species of an urban remnant of a species-rich Themeda triandra grassland in southeastern Australia was quantified and related to abiotic influences. Richness and cover of non-native species were highest at the edges of the remnant and declined to relatively uniform levels within the remnant. Native species richness and cover were lowest at the edge adjoining a roadside but then showed little relation to distance from edge. Roadside edge quadrats were floristically distinct from most other quadrats when ordinated by Detrended Correspondence Analysis. Soil phosphorus was significantly higher at the roadside edge but did not vary within the remnant itself. All other abiotic factors measured (NH₄, NO₃, S, pH and % organic carbon) showed little variation across the remnant. Non-native species richness and cover were strongly correlated with soil phosphorus levels. Native species were negatively correlated with soil phosphorus levels. Canonical Correspondence Analysis identified the perennial non-native grasses of high biomass as species most dependent on high soil nutrient levels. Such species may be resource-limited in undisturbed soils. Three classes of non-native plants have invaded this species-rich grassland: (1) generalist species (> 50 % frequency), mostly therophytes with non-specialized habitat or germination requirements; (2) resource-limited species comprising perennial species of high biomass that are dependent on nutrient increases and/or soil disturbances before they can invade the community and; (3) species of intermediate frequency (1–30 %), of low to high biomass potential, that appear to have non-specialized habitat requirements but are currently limited by seed dispersal, seedling establishment or the current site management. Native species richness and cover are most negatively affected by increases in non-native cover. Declines are largely evident once the non-native cover exceeds 40 %. Widespread, generalist non-native species are numerous in intact sites and will have to be considered a permanent part of the flora of remnant grasslands. Management must aim to minimize increases in cover of any non-native species or the disturbances that favour the establishment of competitive non-native grasses if the native grassland flora is to be conserved in small, fragmented remnants.     

The Role of Tourism and Recreation in the Spread of Non-Native Species: A Systematic Review and Meta-Analysis

Managing the pathways by which non-native species are introduced and spread is considered the most effective way of preventing species invasions. Tourism and outdoor recreation involve the frequent congregation of people, vehicles and vessels from geographically diverse areas. They are therefore perceived to be major pathways for the movement of non-native species, and ones that will become increasingly important with the continued growth of these sectors. However, a global assessment of the relationship between tourism activities and the introduction of non-native species–particularly in freshwater and marine environments–is lacking. We conducted a systematic review and meta-analysis to determine the impact of tourism and outdoor recreation on non-native species in terrestrial, marine and freshwater environments. Our results provide quantitative evidence that the abundance and richness of non-native species are significantly higher in sites where tourist activities take place than in control sites. The pattern was consistent across terrestrial, freshwater and marine environments; across a variety of vectors (e.g. horses, hikers, yachts); and across a range of taxonomic groups. These results highlight the need for widespread biosecurity interventions to prevent the inadvertent introduction of invasive non-native species (INNS) as the tourism and outdoor recreation sectors grow.