Earthworm Invasion as the Driving Force Behind Plant Invasion and Community Change in Northeastern North American Forests

Abstract:  Identification of factors that drive changes in plant community structure and contribute to decline and endangerment of native plant species is essential to the development of appropriate management strategies. Introduced species are assumed to be driving causes of shifts in native plant communities, but unequivocal evidence supporting this view is frequently lacking. We measured native vegetation, non-native earthworm biomass, and leaf-litter volume in 15 forests in the presence and absence of 3 non-native plant species ( Microstegium vimineum, Alliaria petiolata, Berberis thunbergii ) to assess the general impact of non-native plant and earthworm invasions on native plant communities in northeastern United States. Non-native plant cover was positively correlated with total native plant cover and non-native earthworm biomass. Earthworm biomass was negatively associated with cover of native woody and most herbaceous plants and with litter volume. Graminoid cover was positively associated with non-native earthworm biomass and non-native plant cover. These earthworm-associated responses were detected at all sites despite differences in earthworm species and abundance, composition of the native plant community, identity of invasive plant species, and geographic region. These patterns suggest earthworm invasion, rather than non-native plant invasion, is the driving force behind changes in forest plant communities in northeastern North America, including declines in native plant species, and earthworm invasions appear to facilitate plant invasions in these forests. Thus, a focus on management of invasive plant species may be insufficient to protect northeastern forest understory species.     

Impact of Native Plants on Bird and Butterfly Biodiversity in Suburban Landscapes

Abstract:  Managed landscapes in which non-native ornamental plants are favored over native vegetation now dominate the United States, particularly east of the Mississippi River. We measured how landscaping with native plants affects the avian and lepidopteran communities on 6 pairs of suburban properties in southeastern Pennsylvania. One property in each pair was landscaped entirely with native plants and the other exhibited a more conventional suburban mixture of plants—a native canopy with non-native groundcover and shrubs. Vegetation sampling confirmed that total plant cover and plant diversity did not differ between treatments, but non-native plant cover was greater on the conventional sites and native plant cover was greater on the native sites. Several avian (abundance, species richness, biomass, and breeding-bird abundance) and larval lepidopteran (abundance and species richness) community parameters were measured from June 2006 to August 2006. Native properties supported significantly more caterpillars and caterpillar species and significantly greater bird abundance, diversity, species richness, biomass, and breeding pairs of native species. Of particular importance is that bird species of regional conservation concern were 8 times more abundant and significantly more diverse on native properties. In our study area, native landscaping positively influenced the avian and lepidopteran carrying capacity of suburbia and provided a mechanism for reducing biodiversity losses in human-dominated landscapes.     

Wilderness forms and their implications for global environmental policy and conservation

With the intention of securing industry-free land and seascapes, protecting wilderness entered international policy as a formal target for the first time in the zero draft of the Post-2020 Global Biodiversity Framework under the Convention on Biological Diversity. Given this increased prominence in international policy, it is timely to consider the extent to which the construct of wilderness supports global conservation objectives. We evaluated the construct by overlaying recently updated cumulative human pressure maps that offer a global-scale delineation of industry-free land as wilderness with maps of carbon stock, species richness, and ground travel time from urban centers. Wilderness areas took variable forms in relation to carbon stock, species richness, and proximity to urban centers, where 10% of wilderness areas represented high carbon and species richness, 20% low carbon and species richness, and 3% high levels of remoteness (>48 h), carbon, and species richness. Approximately 35% of all remaining wilderness in 2013 was accessible in <24 h of travel time from urban centers. Although the construct of wilderness can be used to secure benefits in specific contexts, its application in conservation must account for contextual and social implications. The diverse characterization of wilderness under a global environmental conservation lens shows that a nuanced framing and application of the construct is needed to improve understanding, communication, and retention of its variable forms as industry-free places.     

Park Management of Exotic Plant Species: Problems and Issues

Abstract: Vegetation management policies in public parks in the United States call for the removal of exotic species to the extent feasible. The underlying goal is to preserve samples of wilderness by restoring plant communities to the "natural state" that existed prior to extensive human influence. With limited budgets, park managers are necessarily selective in targeting exotic species for control. If the focus is on the more readily controlled species, however, park landscapes may gradually become populated by more resistant exotics Further, because plants exhibit some redundancy in ecosystem function, exotic plant species can substitute in part for natives in performing a range of ecosystem functions, including wildlife support and soil binding. Consequently the removal of exotics can result in significant perturbations to certain ecosystem functions during the period of transition to native cover. The individualistic paradigm of plant distribution implies that the impact of exotic plant species on invaded communities will vary. Choosing which species to remove requires careful evaluation of the impact of the removal on ecosystem structure and function. The effective balancing of park management goals for wilderness maintenance and recreational use requires clearer recognition of the adaptive response of ecosystems to invasion and a rethinking of the bases for prioritizing which species are to be removed.     

Thinning and burning result in low-level invasion by nonnative plants but neutral effects on natives.

Many historically fire-adapted forests are now highly susceptible to damage from insects, pathogens, and stand-replacing fires. As a result, managers are employing treatments to reduce fuel loadings and to restore the structure, species, and processes that characterized these forests prior to widespread fire suppression, logging, and grazing. However, the consequences of these activities for understory plant communities are not well understood. We examined the effects of thinning and prescribed fire on plant composition and diversity in Pinus ponderosa forests of eastern Washington (USA). Data on abundance and richness of native and nonnative plants were collected in 70 stands in the Colville, Okanogan, and Wenatchee National Forests. Stands represented one of four treatments: thinning, burning, thinning followed by burning, or control; treatments had been conducted 3-19 years before sampling. Multi-response permutation procedures revealed no significant effect of thinning or burning on understory plant composition. Similarly, there were no significant differences among treatments in cover or richness of native plants. In contrast, nonnative plants showed small, but highly significant, increases in cover and richness in response to both thinning and burning. In the combined treatment, cover of nonnative plants averaged 2% (5% of total plant cover) but did not exceed 7% (16% of total cover) at any site. Cover and richness of nonnative herbs showed small increases with intensity of disturbance and time since treatment. Nonnative plants were significantly less abundant in treated stands than on adjacent roadsides or skid trails, and cover within these potential source areas explained little of the variation in abundance within treated stands. Although thinning and burning may promote invasion of nonnative plants in these forests, our data suggest that their abundance is limited and relatively stable on most sites.     

Estimating non-native plant richness with a species-accumulation model along roads

Monitoring non-native plant richness is important for biodiversity conservation and scientific research. The species-area model (SA model) has been used frequently to estimate the total species richness within a region. However, the conventional SA model may not provide robust estimations of non-native plant richness because the ecological processes associated with the accumulation of exotic and native plants may differ. Because roads strongly dictate the distributions of exotic plants, we propose a species-accumulation model along roads (SR model), rather than an SA model, to estimate the non-native plant richness within a region. Using 270 simulated data sets, we compared the differences in performance between the SR and SA models. A decision tree based on prediction accuracy was created to guide model application, which was validated using field data from 3 national nature reserves in 3 different provinces in China. The SR model significantly outperformed the SA model when non-native species were restricted to the roadsides and the proportion of uncommon exotic species was small. More importantly, the SR model accurately estimated the non-native plant richness in all field sites with an error of <1 species per site. We believe our new model meets the practical need to efficiently and robustly estimate non-native plant richness, which may facilitate effective biodiversity conservations and promote research on non-native plant invasion and vegetation dynamics.     

Rapid assessment of postfire plant invasions in coniferous forests of the western United States.

Fire is a natural part of most forest ecosystems in the western United States, but its effects on nonnative plant invasion have only recently been studied. Also, forest managers are engaging in fuel reduction projects to lessen fire severity, often without considering potential negative ecological consequences such as nonnative plant species introductions. Increased availability of light, nutrients, and bare ground have all been associated with high-severity fires and fuel treatments and are known to aid in the establishment of nonnative plant species. We use vegetation and environmental data collected after wildfires at seven sites in coniferous forests in the western United States to study responses of nonnative plants to wildfire. We compared burned vs. unburned plots and plots treated with mechanical thinning and/or prescribed burning vs. untreated plots for nonnative plant species richness and cover and used correlation analyses to infer the effect of abiotic site conditions on invasibility. Wildfire was responsible for significant increases in nonnative species richness and cover, and a significant decrease in native cover. Mechanical thinning and prescribed fire fuel treatments were associated with significant changes in plant species composition at some sites. Treatment effects across sites were minimal and inconclusive due to significant site and site x treatment interaction effects caused by variation between sites including differences in treatment and fire severities and initial conditions (e.g., nonnative species sources). We used canonical correspondence analysis (CCA) to determine what combinations of environmental variables best explained patterns of nonnative plant species richness and cover. Variables related to fire severity, soil nutrients, and elevation explained most of the variation in species composition. Nonnative species were generally associated with sites with higher fire severity, elevation, percentage of bare ground, and lower soil nutrient levels and lower canopy cover. Early assessments of postfire stand conditions can guide rapid responses to nonnative plant invasions.     

Declines in Woodland Salamander Abundance Associated with Non-Native Earthworm and Plant Invasions

Abstract:  Factors that negatively affect the quality of wildlife habitat are a major concern for conservation. Non-native species invasions, in particular, are perceived as a global threat to the quality of wildlife habitat. Recent evidence indicates that some changes to understory plant communities in northern temperate forests of North America, including invasions by 3 non-native plant species, are facilitated by non-native earthworm invasion. Furthermore, non-native earthworm invasions cause a reduction in leaf litter on the forest floor, and the loss of forest leaf litter is commonly associated with declines in forest fauna, including amphibians. We conducted a mark-recapture study of woodland salamander abundance across plant invasion fronts at 10 sites to determine whether earthworm or plant invasions were associated with reduced salamander abundance. Salamander abundance declined exponentially with decreasing leaf litter volume. There was no significant relationship between invasive plant cover and salamander abundance, independent of the effects of leaf litter loss due to earthworm invasion. An analysis of selected salamander prey abundance (excluding earthworms) at 4 sites showed that prey abundance declined with declining leaf litter. The loss of leaf litter layers due to non-native earthworm invasions appears to be negatively affecting woodland salamander abundance, in part, because of declines in the abundance of small arthropods that are a stable resource for salamanders. Our results demonstrate that earthworm invasions pose a significant threat to woodland amphibian fauna in the northeastern United States, and that plant invasions are symptomatic of degraded amphibian habitat but are not necessarily drivers of habitat degradation.     

Aquatic plant community invasibility and scale-dependent patterns in native and invasive species richness

Invasive species richness often is negatively correlated with native species richness at the small spatial scale of sampling plots, but positively correlated in larger areas. The pattern at small scales has been interpreted as evidence that native plants can competitively exclude invasive species. Large-scale patterns have been understood to result from environmental heterogeneity, among other causes. We investigated species richness patterns among submerged and floating-leaved aquatic plants (87 native species and eight invasives) in 103 temperate lakes in Connecticut (northeastern USA) and found neither a consistently negative relationship at small (3-m2) scales, nor a positive relationship at large scales. Native species richness at sampling locations was uncorrelated with invasive species richness in 37 of the 60 lakes where invasive plants occurred; richness was negatively correlated in 16 lakes and positively correlated in seven. No correlation between native and invasive species richness was found at larger spatial scales (whole lakes and counties). Increases in richness with area were uncorrelated with abiotic heterogeneity. Logistic regression showed that the probability of occurrence of five invasive species increased in sampling locations (3 m2, n = 2980 samples) where native plants occurred, indicating that native plant species richness provided no resistance against invasion. However, the probability of three invasive species' occurrence declined as native plant density increased, indicating that density, if not species richness, provided some resistance with these species. Density had no effect on occurrence of three other invasive species. Based on these results, native species may resist invasion at small spatial scales only in communities where density is high (i.e., in communities where competition among individuals contributes to community structure). Most hydrophyte communities, however, appear to be maintained in a nonequilibrial condition by stress and/or disturbance. Therefore, most aquatic plant communities in temperate lakes are likely to be vulnerable to invasion.     

Effects of Earthworm Invasion on Plant Species Richness in Northern Hardwood Forests

Abstract:  The invasion of non-native earthworms ( Lumbricus spp.) into a small number of intensively studied stands of northern hardwood forest has been linked to declines in plant diversity and the local extirpation of one threatened species. It is unknown, however, whether these changes have occurred across larger regions of hardwood forests, which plant species are most vulnerable, or with which earthworm species such changes are associated most closely. To address these issues we conducted a regional survey in the Chippewa and Chequamegon national forests in Minnesota and Wisconsin (U.S.A.), respectively. We sampled earthworms, soils, and vegetation, examined deer browse in 20 mature, sugar-maple-dominated forest stands in each national forest, and analyzed the relationship between invasive earthworms and vascular plant species richness and composition. Invasion by Lumbricus was a strong indicator of reduced plant richness in both national forests. The mass of Lumbricus juveniles was significantly and negatively related to plant-species richness in both forests. In addition, Lumbricus was a significant factor affecting plant richness in a full model that included multiple variables. In the Chequamegon National Forest earthworm mass was associated with higher sedge cover and lower cover of sugar maple seedlings and several forb species. The trends were similar but not as pronounced in Chippewa, perhaps due to lower deer densities and different earthworm species composition. Our results provide regional evidence that invasion by Lumbricus species may be an important mechanism in reduced plant-species richness and changes in plant communities in mature forests dominated by sugar maples.     

Effects of vegetation cover, presence of a native ant species, and human disturbance on colonization by Argentine ants

The spread of non-native invasive species is affected by human activity, vegetation cover, weather, and interaction with native species. We analyzed data from a 17-year study of the distribution of the non-native Argentine ant (Linepithema humile) and the native winter ant (Prenolepis imparis) in a preserve in northern California (U.S.A.). We conducted logistic regressions and used model selection to determine whether the following variables were associated with changes in the distribution of each species: presence of conspecifics at neighboring sites, distance to development (e.g., roads, buildings, and landscaped areas), proportion of vegetation cover taller than 0.75 m, elevation, distance to water, presence of both species at a site, temperature, and rainfall. Argentine ants colonized unoccupied sites from neighboring sites, but the probability of appearance and persistence decreased as distance to development, vegetation cover, and elevation increased. Winter ants appeared and persisted in sites with relatively high vegetation cover (i.e., highly shaded sites). Presence of the 2 species was negatively associated in sites with high vegetation cover (more winter ants) and sites near development (more Argentine ants). Probability of colonization of Argentine ants decreased where winter ants were most persistent. At sites near development within the preserve, abundant Argentine ant populations may be excluding winter ants. The high abundance of Argentine ants at these sites may be due to immigration from suburban areas outside the preserve, which are high-quality habitat for Argentine ants. In the interior of the preserve, distance from development, low-quality habitat, and interaction with winter ants may in combination exclude Argentine ants. Interactions among the variables we examined were associated with low probabilities of Argentine ant colonization in the preserve.     

High richness and dense seeding enhance grassland restoration establishment but have little effect on drought response

Restorations commonly utilize seed addition to formerly arable lands where the development of native plant communities is severely dispersal limited. However, variation in seed addition practices may profoundly affect restoration outcomes. Theory and observations predict that species-rich seed mixtures and seeding at high densities should enhance native plant community establishment, minimize exotic species cover, and may promote resistance and resilience to, and recovery from, environmental perturbations. We studied the post-seeding establishment of native plant communities in large grassland restoration plots, which were sown at two densities crossed with two levels of species richness on formerly arable land in Nebraska, USA, and their responses to drought. To evaluate drought resistance, recovery, and resilience of restored plant communities, we erected rainfall manipulation structures and tracked the response of seeded species cover and total plant biomass during experimental drought relative to controls and in the post-drought growing season. High seed richness and high-density seeding treatments resulted in greater richness and cover of native, seeded species per 0.5 m2 compared to low-richness and low-density treatments. Cover differences in response to seed mixture richness were driven by native forbs. Richness and cover of exotic species were lowest in high-richness and high-density treatments. We found little evidence of differential drought resistance, recovery, and resilience among seeding treatments. Increases in exotic species across years were restricted to drought subplots, and were not affected by seeding treatments. Grassland restoration was generally enhanced and exotic cover reduced both by the use of high-richness seed mixtures and high-density seeding. Given the lack of restoration treatment effects on the resistance, recovery, or resilience of seeded species exposed to drought, and the increases in exotic species following drought, other forms of active management may be needed to produce restored plant communities that are robust to climate change.     

Fuel breaks affect nonnative species abundance in Californian plant communities.

We evaluated the abundance of nonnative plants on fuel breaks and in adjacent untreated areas to determine if fuel treatments promote the invasion of nonnative plant species. Understanding the relationship between fuel treatments and nonnative plants is becoming increasingly important as federal and state agencies are currently implementing large fuel treatment programs throughout the United States to reduce the threat of wildland fire. Our study included 24 fuel breaks located across the State of California. We found that nonnative plant abundance was over 200% higher on fuel breaks than in adjacent wildland areas. Relative nonnative cover was greater on fuel breaks constructed by bulldozers (28%) than on fuel breaks constructed by other methods (7%). Canopy cover, litter cover, and duff depth also were significantly lower on fuel breaks constructed by bulldozers, and these fuel breaks had significantly more exposed bare ground than other types of fuel breaks. There was a significant decline in relative nonnative cover with increasing distance from the fuel break, particularly in areas that had experienced more numerous fires during the past 50 years, and in areas that had been grazed. These data suggest that fuel breaks could provide establishment sites for nonnative plants, and that nonnatives may invade surrounding areas, especially after disturbances such as fire or grazing. Fuel break construction and maintenance methods that leave some overstory canopy and minimize exposure of bare ground may be less likely to promote nonnative plants.     

Spatial heterogeneity influences native and nonnative plant species richness

Spatial heterogeneity may have differential effects on the distribution of native and nonnative plant species richness. We examined the effects of spatial heterogeneity on native and nonnative plant species richness distributions in the central part of Rocky Mountain National Park, Colorado, USA. Spatial heterogeneity around vegetation plots was characterized using landscape metrics, environmental/topographic variables (slope, aspect, elevation, and distance from stream or river), and soil variables (nitrogen, clay, and sand). The landscape metrics represented five components of landscape heterogeneity and were measured at four spatial extents (within varying radii of 120, 240, 480, and 960 m) using the FRAGSTATS landscape pattern analysis program. Akaike's Information Criterion adjusted for small sample size (AICc) was used to select the best models from a set of multiple linear regression models developed for native and nonnative plant species richness at four spatial extents and three levels of ecological hierarchy (i.e., landscape, land cover, and community). Both native and nonnative plant species richness were positively correlated with edge density, Simpson's diversity index and interspersion/juxtaposition index, and were negatively correlated with mean patch size. The amount of variation explained at four spatial extents and three hierarchical levels ranged from 30% to 70%. At the landscape level, the best models explained 43% of the variation in native plant species richness and 70% of the variation in nonnative plant species richness (240-m extent). In general, the amount of variation explained was always higher for nonnative plant species richness, and the inclusion of landscape metrics always significantly improved the models. The best models explained 66% of the variation in nonnative plant species richness for both the conifer land cover type and lodgepole pine community. The relative influence of the components of spatial heterogeneity differed for native and nonnative plant species richness and varied with the spatial extent of analysis and levels of ecological hierarchy. The study offers an approach to quantify spatial heterogeneity to improve models of plant biodiversity. The results demonstrate that ecologists must recognize the importance of spatial heterogeneity in managing native and nonnative plant species.     

Susceptibility of a Northern Hardwood Forest to Exotic Earthworm Invasion

Abstract:  Numerous exotic earthworm species are colonizing northern hardwood forests of North America, where no native earthworms exist. Upon invasion, earthworms have been shown to alter the surface soil environment and plant populations and communities. We sought to identify land-use factors in the Ottawa National Forest (ONF), Michigan (U.S.A.), that contribute to earthworm invasion in forest dominated by sugar maple ( Acer saccharum Marsh.) so that the susceptibility to additional colonization could be evaluated. We sampled earthworm communities in Sylvania Wilderness Area, a unique old-growth hardwood forest, and nonwilderness sites influenced by recreational fishing, recent timber harvesting, or roads. All the nonwilderness sites contained one to five species of exotic earthworms. In contrast, only 50% of wilderness sites contained exotic earthworms, all of a single species. Nonwilderness sites also had thinner litter and duff layers, higher soil C and N content, and higher nitrogen mineralization potentials than Sylvania sites. Two central differences between Sylvania and nonwilderness sites were that all nonwilderness sites were in close contact with roads and had a history of timber harvest, whereas these factors were not present in Sylvania Wilderness Area. Using average rates of colonization, we constructed two geographic information system models to estimate the percentage of sugar maple on the ONF falling within a theoretical 100-year invasion distance of roads and of second-growth sugar maple as relative indices of susceptibility to invasion. Both models indicated high susceptibility to invasion, with 91.7% and 98.9% of sugar maple habitat falling within a theoretical 100-year invasion distance of roads or historical harvests, respectively.     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

Protected‐Area Boundaries as Filters of Plant Invasions

Abstract: Human land uses surrounding protected areas provide propagules for colonization of these areas by non‐native species, and corridors between protected‐area networks and drainage systems of rivers provide pathways for long‐distance dispersal of non‐native species. Nevertheless, the influence of protected‐area boundaries on colonization of protected areas by invasive non‐native species is unknown. We drew on a spatially explicit data set of more than 27,000 non‐native plant presence records for South Africa's Kruger National Park to examine the role of boundaries in preventing colonization of protected areas by non‐native species. The number of records of non‐native invasive plants declined rapidly beyond 1500 m inside the park; thus, we believe that the park boundary limited the spread of non‐native plants. The number of non‐native invasive plants inside the park was a function of the amount of water runoff, density of major roads, and the presence of natural vegetation outside the park. Of the types of human‐induced disturbance, only the density of major roads outside the protected area significantly increased the number of non‐native plant records. Our findings suggest that the probability of incursion of invasive plants into protected areas can be quantified reliably.     

Anthropogenic Correlates of Species Richness in Southeastern Ontario Wetlands

We examined the relationship between the richness of four different wetland taxa (birds, mammals, herptiles, and plants) in 30 southeastern Ontario, Canada wetlands and two anthropogenic factors: road construction and forest removal/conversion on adjacent lands. Data were obtained from two sources: road densities and forest cover from 1:50,000 Government of Canada topographic maps and species lists and wetland areas from Ontario Ministry of Natural Resources wetland evaluation reports. Multiple regression analysis was used to model the relationships between species richness and wetland area, road density, and forest cover. Our results show a strong positive relationship between wetland area and species richness for all taxa. The species richness of all taxa except mammals was negatively correlated with the density of paved roads on lands up to 2 km from the wetland. Furthermore, both herptile and mammal species richness showed a strong positive correlation with the proportion of forest cover on lands within 2 km. These results provide evidence that at the landscape level, road construction and forest removal on adjacent lands pose significant risks to wetland biodiversity. Furthermore, they suggest that most existing wetland policies, which focus almost exclusively on activities within the wetland itself and/or a narrow buffer zone around the wetland perimeter, are unlikely to provide adequate protection for wetland biodiversity.     

Changing Species Richness and Composition in Canadian National Parks

Abstract: Canada's national parks and their surrounding areas differ markedly in size, climate, vegetation, and extent of human development. We tested the extent to which total species richness, native species richness, and the number of extirpations and introductions of terrestrial vertebrates were correlated with each of these factors. To do this, we used surveys of park fauna from the present and from the time of park establishment. Richness, extirpations, and introductions were all strongly related to climate. After we controlled for climate, smaller parks had higher rates of species loss than larger parks. Land-use patterns (forest cover and fragmentation, roads, etc.) within parks were strongly correlated with land use in the regions surrounding the parks, showing that parks have not been isolated from regional development. Richness and extirpations within parks were generally more strongly related to regional characteristics than to the characteristics of the parks themselves. Species richness and numbers of introduced species were higher in parks found in landscapes with greater fragmentation. Frequencies of extirpations were less clearly related to human-influenced habitat characteristics. Introductions and extinctions most often involved game species or species directly associated with human activities. There is little evidence of subtle ecological effects being responsible for species loss. Our results suggest that management should focus on direct human interventions, such as hunting, introduction of game species, and habitat fragmentation, in parks and surrounding areas.     

Worldwide effects of non-native species on species–area relationships

Non-native species have invaded most parts of the world, and the invasion process is expected to continue and accelerate. Because many invading non-native species are likely to become permanent inhabitants, future consideration of species-area relationships (SARs) should account for non-native species, either separately or jointly with native species. If non-native species occupy unused niches and space in invaded areas and extinction rate of native species remains low (especially for plants), the resultant SARs (with both native and non-native species) will likely be stronger. We used published and newly compiled data (35 data sets worldwide) to examine how species invasions affect SARs across selected taxonomic groups and diverse ecosystems around the world. We first examined the SARs for native, non-native, and all species. We then investigated with linear regression analyses and paired or unpaired t tests how degree of invasion (proportion of non-native species) affected postinvasion SARs. Postinvasion SARs for all species (native plus non-native) became significantly stronger as degree of invasion increased (r² = 0.31, p = 0.0006), thus, reshaping SARs worldwide. Overall, native species still showed stronger and less variable SARs. Also, slopes for native species were steeper than for non-native species (0.298 vs. 0.153). There were some differences among non-native taxonomic groups in filling new niches (especially for birds) and between islands and mainland ecosystems. We also found evidence that invasions may increase equilibrial diversity. Study of such changing species–area curves may help determine the probability of future invasions and have practical implications for conservation.