Spatial patch occupancy patterns of the Lower Keys marsh rabbit

Reliable estimates of presence or absence of a species can provide substantial information on management questions related to distribution and habitat use but should incorporate the probability of detection to reduce bias. We surveyed for the endangered Lower Keys marsh rabbit (Sylvilagus palustris hefneri) in habitat patches on 5 Florida Key islands, USA, to estimate occupancy and detection probabilities. We derived detection probabilities using spatial replication of plots and evaluated hypotheses that patch location (coastal or interior) and patch size influence occupancy and detection. Results demonstrate that detection probability, given rabbits were present, was <0.5 and suggest that naïve estimates (i.e., estimates without consideration of imperfect detection) of patch occupancy are negatively biased. We found that patch size and location influenced probability of occupancy but not detection. Our findings will be used by Refuge managers to evaluate population trends of Lower Keys marsh rabbits from historical data and to guide management decisions for species recovery. The sampling and analytical methods we used may be useful for researchers and managers of other endangered lagomorphs and cryptic or fossorial animals occupying diverse habitats. © 2011 The Wildlife Society.     

Population Structure of the Lower Keys Marsh Rabbit as Determined by Mitochondrial DNA Analysis

ABSTRACT We used nucleotide sequence data from a mitochondrial DNA fragment to characterize variation within the endangered Lower Keys marsh rabbit (Sylvilagus palustris hefneri). We observed 5 unique mitochondrial haplotypes across different sampling sites in the Lower Florida Keys, USA. Based on the frequency of these haplotypes at different geographic locations and relationships among haplotypes, we observed 2 distinct clades or groups of sampling sites (western and eastern clades). These 2 groups showed low levels of gene flow. Regardless of their origin, marsh rabbits from the Lower Florida Keys can be separated into 2 genetically distinct management units, which should be considered prior to implementation of translocations as a means of offsetting recent population declines.     

Occupancy of yellow‐billed and Pacific loons: evidence for interspecific competition and habitat mediated co‐occurrence

Interspecific competition is an important process structuring ecological communities, however, it is difficult to observe in nature. We used an occupancy modelling approach to evaluate evidence of competition between yellow‐billed (Gavia adamsii) and Pacific (G. pacifica) loons for nesting lakes on the Arctic Coastal Plain of Alaska. With multiple years of data and survey platforms, we estimated dynamic occupancy states (e.g. rates of colonization or extinction from individual lakes) and controlled for detection differences among aircraft platforms and ground survey crews. Results indicated that yellow‐billed loons were strong competitors and negatively influenced the occupancy of Pacific loons by excluding them from potential breeding lakes. Pacific loon occupancy was conditional on the presence of yellow‐billed loons, with Pacific loons having almost a tenfold decrease in occupancy probability when yellow‐billed loons were present and a threefold decrease in colonization probability when yellow‐billed loons were present in the current or previous year. Yellow‐billed and Pacific loons co‐occurred less than expected by chance except on very large lakes or lakes with convoluted shorelines; variables which may decrease the cost of maintaining a territory in the presence of the other species. These results imply the existence of interspecific competition between yellow‐billed and Pacific loons for nesting lakes; however, habitat characteristics which facilitate visual and spatial separation of territories can reduce competitive interactions and promote species co‐occurrence.     

Human disturbance and stage‐specific habitat requirements influence snowy plover site occupancy during the breeding season

Habitat use has important consequences for avian reproductive success and survival. In coastal areas with recreational activity, human disturbance may limit use of otherwise suitable habitat. Snowy plovers Charadrius nivosus have a patchy breeding distribution along the coastal areas on the Florida Panhandle, USA. Our goal was to determine the relative effects of seasonal human disturbance and habitat requirements on snowy plover habitat use. We surveyed 303 sites for snowy plovers, human disturbance, and habitat features between January and July 2009 and 2010. We made multiple visits during three different sampling periods that corresponded to snowy plover breeding: pre‐breeding, incubation, and brood‐rearing and used multi‐season occupancy models to examine whether human disturbance, habitat features, or both influenced site occupancy, colonization (probability of transition from an unoccupied site to an occupied site), and extinction (probability of transition from an occupied site to an unoccupied site). Snowy plover site occupancy and colonization was negatively associated with human disturbance and site extinction was positively associated with human disturbance. Interdune vegetation had a negative effect on occupancy and colonization, indicating that plovers were less likely to use areas with uniform, dense vegetation among dunes. Also, dune shape, beach debris, and access to low‐energy foraging areas influenced site occupancy, colonization, and extinction. Plovers used habitat based on beach characteristics that provided stage‐specific resource needs; however, human disturbance was the strongest predictor of site occupancy. In addition, vegetation plantings used to enhance dune rehabilitation may negatively impact plover site occupancy. Management actions that decrease human disturbance, such as symbolic fencing and signage, may increase the amount of breeding habitat available to snowy plovers on the Florida Panhandle and in other areas with high human activity. The specific areas that require this protection may vary across snowy plover life history stages.     

Testing metapopulation concepts: effects of patch characteristics and neighborhood occupancy on the dynamics of an endangered lagomorph

Metapopulation ecology is a field that is richer in theory than in empirical results. Many existing empirical studies use an incidence function approach based on spatial patterns and key assumptions about extinction and colonization rates. Here we recast these assumptions as hypotheses to be tested using 18 years of historic detection survey data combined with four years of data from a new monitoring program for the Lower Keys marsh rabbit. We developed a new model to estimate probabilities of local extinction and colonization in the presence of nondetection, while accounting for estimated occupancy levels of neighboring patches. We used model selection to identify important drivers of population turnover and estimate the effective neighborhood size for this system. Several key relationships related to patch size and isolation that are often assumed in metapopulation models were supported: patch size was negatively related to the probability of extinction and positively related to colonization, and estimated occupancy of neighboring patches was positively related to colonization and negatively related to extinction probabilities. This latter relationship suggested the existence of rescue effects. In our study system, we inferred that coastal patches experienced higher probabilities of extinction and colonization than interior patches. Interior patches exhibited higher occupancy probabilities and may serve as refugia, permitting colonization of coastal patches following disturbances such as hurricanes and storm surges. Our modeling approach should be useful for incorporating neighbor occupancy into future metapopulation analyses and in dealing with other historic occupancy surveys that may not include the recommended levels of sampling replication.     

Landscape Ecology of Eastern Spotted Skunks in Habitats Restored for Red‐Cockaded Woodpeckers

Although examples are rare, conflicts between species of conservation concern can result from habitat restoration that modifies habitat to benefit a single taxon. A forest restoration program designed to enhance habitat for endangered red‐cockaded woodpeckers (Picoides borealis) may be reducing available habitat for the eastern spotted skunk (Spilogale putorius), a forest‐adapted sympatric species of conservation concern that occurs in the Ouachita National Forest, Arkansas, U.S.A. At small scales, eastern spotted skunks select early successional forest with structural diversity, whereas red‐cockaded woodpeckers prefer mature pine (Pinus spp.) habitat. We surveyed for eastern spotted skunks at 50 managed forest stands, modeled occupancy as a function of landscape‐level habitat factors to examine how features of restoration practices influenced occurrence, and compared known occupied forest stands to those where active red‐cockaded woodpecker nesting clusters were located. The most‐supported occupancy models contained forest stand age (negatively associated) and size (positively associated); suggesting eastern spotted skunks primarily occupy large patches of habitat with dense understory and overhead cover. Red‐cockaded woodpecker nesting clusters were located in smaller and older forest stands. These results suggest that increased overhead cover, which can reduce risk of avian predation, enhances occupancy by small forest carnivores such as eastern spotted skunks. Management activities that increase forest stand rotation length reduce the availability of young dense forest. The practice may enhance the value of habitat for red‐cockaded woodpeckers, but may reduce the occurrence of eastern spotted skunks. Implementing plans that consider critical habitat and extinction risks for multiple species may reduce such conservation conflict.     

Evaluating nest supplementation as a recovery strategy for the endangered rodents of the Florida Keys

The Key Largo woodrat (Neotoma floridana smalli) and Key Largo cotton mouse (Peromyscus gossypinus allapaticola) are federally endangered subspecies endemic to the tropical hardwood hammocks of Key Largo, Florida. Woodrats are considered generalists in habitat and diet, yet a steady decline in natural stick nests and capture rates over the past several decades suggests that they are limited by the availability of nesting habitat due to habitat loss and fragmentation. The more specialized Key Largo cotton mouse appears to rely on old growth hammock, a habitat type that is rare following past land clearing. In 2004, the U.S. Fish and Wildlife Service started building supplemental nest structures to restore habitat quality and connectivity for these endangered rodents, but nest use requires evaluation. We used camera traps and occupancy models to evaluate the factors influencing woodrat and cotton mouse use of the supplemental nests. We detected woodrats at 65 and cotton mice at 175 of 284 sampled nest structures, with co‐occurrence at 38 nests. Woodrat nest use followed a gradient from low nest use in the north to high nest use in the south, which might relate to the proximity of free‐ranging domestic cat (Felis catus) colonies in residential developments. Cotton mouse nest use, however, was related positively to mature hammock and related negatively to disturbed areas (e.g. scarified lands). The two species occurred independently of each other. Stick‐stacking behavior was observed at supplemental nests and, although it was correlated with detection of woodrats, it was not a strong predictor of their occurrence. We suggest that nest supplementation can be an important tool for species recovery as habitat quality continues to improve with succession.     

Evidence for a recent genetic bottleneck in the endangered Florida Keys silver rice rat (<Emphasis Type="Italic">Oryzomys argentatus</Emphasis>) revealed by microsatellite DNA analyses

Low levels of genetic variation are thought to contribute significantly to the higher extinction rates of endemic island populations compared to their mainland counterparts. We used six microsatellite loci to compare the genetic structure of the endangered silver rice rat (Oryzomys argentatus) population in Saddlebunch Key, Florida to the mainland population of the closely related marsh rice rat (Oryzomys palustris natator) in Everglades National Park. Allelic richness and gene diversity are significantly lower in Saddlebunch Key than in the larger mainland population, and the two populations are significantly differentiated as measured by both F-statistics and Bayesian clustering methods. These findings support the classification of the Keys population as a “distinct vertebrate population” by the U.S. Fish and Wildlife Service. Current gene diversity (H _E) is higher than expected under mutation-drift equilibrium in Saddlebunch Key, indicating a genetic bottleneck. The Keys population also exhibits a mode shift in its allele frequency distribution which suggests a very recent bottleneck has occurred and is consistent with reports of recent population declines. Although habitat loss and exotic species pose a more immediate and serious threat to silver rice rats, the continued loss of genetic variation may contribute to their long-term extinction risk due to inbreeding or by lowering the population’s ability to adapt to future environmental changes. The protection of habitat and the removal of introduced predators and competitors may help increase the population size of silver rice rats and lower their risk of extinction, both from a demographic and a genetic perspective.     

Occupancy, colonization and extinction patterns of rabbit populations: implications for Iberian lynx conservation

In Mediterranean ecosystems, rabbits are a key prey species for many predators, such as the Iberian lynx, which is threatened with extinction and has gone extinct locally in several regions of its historical distribution range. One of these regions is Serra da Malcata Nature Reserve, Portugal, which is also currently proposed as a potential site for reintroduction. We intended to investigate annual variation, potential time trends and the effects of management practices on the rabbit population in Serra da Malcata as a model for future potential reintroduction areas. The rabbit population was monitored over 12?years (from 1997 to 2009) by counting latrines along linear transects. These data were used to estimate rabbit occupancy, colonization and extinction patterns using a likelihood-based method including habitat, population and topographic covariate effects. Our results suggest that initial occupancy, when management practices were absent, was driven by the presence of Erica spp. and Cistus ladanifer shrubs and by distance to summits. Site colonization was positively influenced by the presence of edges between shrubs and pastureland and by patterns of rabbit distribution in the previous sampling season. On the other hand, local extinction was negatively influenced by edges. We conclude that the increase in rabbit occupancy and local colonization patterns was clearly associated with management actions (particularly, the creation of pasturelands), although the recovery of the species was noticeably limited by previous patterns of spatial distribution.     

Spatio‐temporal variation of biotic factors underpins contemporary range dynamics of congeners

Species’ ranges are complex often exhibiting multidirectional shifts over space and time. Despite the strong fingerprint of recent historical climate change on species’ distributions, biotic factors such as loss of vegetative habitat and the presence of potential competitors constitute important yet often overlooked drivers of range dynamics. Furthermore, short‐term changes in environmental conditions can influence the underlying processes of local extinction and local colonization that drive range shifts, yet are rarely considered at broad scales. We used dynamic state‐space occupancy models to test multiple hypotheses of the relative importance of major drivers of range shifts of Golden‐winged Warblers (Vermivora chrysoptera) and Blue‐winged Warblers (V. cyanoptera) between 1983 and 2012 across North America: warming temperatures; habitat changes; and occurrence of congeneric species, used here as proxy for biotic interactions. Dynamic occupancies for both species were most influenced by spatial relative to temporal variation in temperature and habitat. However, temporal variation in temperature anomalies and biotic interactions remained important. The two biotic factors considered, habitat change and biotic interactions, had the largest relative effect on estimated extinction rates followed by abiotic temperature anomalies. For the Golden‐winged Warbler, the predicted presence of the Blue‐winged Warbler, a hypothesized competitor, most influenced extinction probabilities, contributing to evidence supporting its role in site‐level species replacement. Given the overall importance of biotic factors on range‐wide dynamic occupancies, their consideration alongside abiotic factors should not be overlooked. Our results suggest that warming compounds the negative effect of habitat loss emphasizing species’ need for habitat to adapt to a changing climate. Notably, even closely related species exhibited individual responses to abiotic and biotic factors considered.     

Occupancy patterns of an apex avian predator across a forest landscape

Apex predators are integral parts of every ecosystem, having top‐down roles in food web maintenance. Understanding the environmental and habitat characteristics associated with predator occurrence is paramount to conservation efforts. However, detecting top order predators can be difficult due to small population sizes and cryptic behaviour. The endangered Tasmanian masked owl (Tyto novaehollandiae castanops) is a nocturnal predator with a distribution understood to be associated with high mature forest cover at broad scales. With the aim to gather monitoring data to inform future conservation effort, we trialled an occupancy survey design to model masked owl occurrence across ~800 km² in the Tasmanian Southern Forests. We conducted 662 visits to assess masked owl occupancy at 160 sites during July–September 2018. Masked owl site occupancy was 12%, and estimated detectability was 0.26 (±0.06 SE). Cumulative detection probability of masked owls over four visits was 0.7. Occupancy modelling suggested owls were more likely to be detected when mean prey count was higher. However, low detection rates hindered the development of confident occupancy predictions. To inform effective conservation of the endangered Tasmanian masked owl, there is a need to develop novel survey techniques that better account for the ecology of this rare, wide‐ranging and cryptic predator. We discuss the potential to combine novel census approaches that exploit different aspects of masked owl ecology to obtain more robust and detailed data.     

Feeling the pressure at home: Predator activity at the burrow entrance of an endangered arid‐zone skink

Habitat modification and invasive species are among the most important contemporary drivers of biodiversity loss. These two threatening processes are often studied independently and few studies have focused on how they interact to influence species declines. Here we assess the predation pressure placed on the threatened great desert skink (Liopholis kintorei) and how this interacts with fire‐induced habitat modifications. We collected daily track data of potential predators for 1 month at 30 great desert skink burrow‐systems where vegetation cover varied significantly after experimental burns. We used these data to evaluate potential predation pressure at the burrow‐system and assess whether fire influenced predator pressure. We supplemented this analysis by documenting predation via the inspection of large mammalian predator scats collected from great desert skink habitat. The level of feral cat activity at a burrow‐system entrance was significantly higher than that of any other potential predator, however fire had no effect on the visitation rates of feral cats, dingoes or large snakes to great desert skink burrow‐systems. The remains of great desert skink were found significantly more frequently in feral cat scats, compared to fox and dingo scats. We provide the first direct evidence that feral cats are a significant predator for great desert skink, thus supporting the hypothesis that feral cat predation is a key threatening process. Feral cat activity was not influenced by small‐scale experimental burns, however, this does not preclude an effect of larger scale fires and we recommend further research exploring this possible interaction.     

Experimental predator removal causes rapid salt marsh die‐off

Salt marsh habitat loss to vegetation die‐offs has accelerated throughout the western Atlantic in the last four decades. Recent studies have suggested that eutrophication, pollution and/or disease may contribute to the loss of marsh habitat. In light of recent evidence that predators are important determinants of marsh health in New England, we performed a total predator exclusion experiment. Here, we provide the first experimental evidence that predator depletion can cause salt marsh die‐off by releasing the herbivorous crab Sesarma reticulatum from predator control. Excluding predators from a marsh ecosystem for a single growing season resulted in a >100% increase in herbivory and a >150% increase in unvegetated bare space compared to plots with predators. Our results confirm that marshes in this region face multiple, potentially synergistic threats.     

Impacts of a half century of sea‐level rise and development on an endangered mammal

The extraordinary growth of human populations and development in coastal areas over the last half century has eliminated and degraded coastal habitats and threatened the persistence of associated wildlife. Moreover, human‐induced sea‐level rise (SLR) is projected to further eliminate and alter the same coastal ecosystems, especially low‐lying regions. Whereas habitat loss and wildlife population declines from development are well documented, contemporary SLR has not yet been implicated in declines of coastal faunal populations. In addition, the projection of severe synergistic impacts from the combination of development and SLR is well described, yet the scientific literature offers little empirical evidence of the influence of these forces on coastal wildlife. Analysis of aerial photographs from 1959 to 2006 provided evidence of a 64% net loss of the endangered Lower Keys marsh rabbit's (Sylvilagus palustris hefneri; LKMR) habitat, the majority due to SLR (>48%). Furthermore, there was a strong negative relationship between the proportion of development per island and the amount of new habitat formed. Islands with modest development (less than 8% of land area) saw formation of new areas of marsh vegetation suitable for rabbits, whereas islands with 8% or more of their lands developed between 1959 and 2006 saw little to no addition of LKMR habitat. Only 8% of habitat loss was directly due to conversion to impervious surfaces, indicating that the greatest threats from development were indirect, including blocking of the inland migration of habitat triggered by SLR. Our results were consistent with an ongoing squeeze of coastal ecosystems between rising seas and development as a threat to LKMR habitat, which raises concern for a wide variety of coastal species. Our results provide evidence that SLR has become a contemporary conservation concern, one that is exacerbated by development, and expected to increase in magnitude as ocean waters continue to rise.     

Scale‐dependent strategies for coexistence of mesocarnivores in human‐dominated landscapes

Identifying factors influencing the distribution of and interactions within carnivore communities is important for understanding how they are affected by human activities. Species differ in their ability to adapt to humans depending on their degree of specialization in habitat use and feeding habits. This results in asymmetric changes in the ecology of co‐occurring species that can influence their interactions. We investigated whether human infrastructures and free‐ranging domestic dogs (a species typically associated with humans) influenced the co‐occurrence and habitat use of mesocarnivores in a landscape of high human population density in Maharashtra, India. We used 40 camera trap locations during 233 trapping nights and used Bayesian co‐occurrence occupancy models to investigate the habitat use and coexistence of species at different spatial scales. Additionally, we investigated their temporal overlap in space use. Indian foxes altered their habitat use both spatially and temporally in order to avoid free‐ranging domestic dogs and other larger competitors. The use of human infrastructure by jackals and jungle cats was limited by the presence of dogs. Our results illustrate how habitat use of smaller carnivore species changes both spatially and temporally in order to avoid larger competitors. We also show that the presence of species associated with humans mediates the influence of human infrastructures on the habitat use of mesocarnivores. We highlight the importance of acknowledging the potential impact of urbanization not only on single species, but also on the interactions within the community.     

Occupancy versus colonization–extinction models for projecting population trends at different spatial scales

Understanding spatiotemporal population trends and their drivers is a key aim in population ecology. We further need to be able to predict how the dynamics and sizes of populations are affected in the long term by changing landscapes and climate. However, predictions of future population trends are sensitive to a range of modeling assumptions. Deadwood‐dependent fungi are an excellent system for testing the performance of different predictive models of sessile species as these species have different rarity and spatial population dynamics, the populations are structured at different spatial scales, and they utilize distinct substrates. We tested how the projected large‐scale occupancies of species with differing landscape‐scale occupancies are affected over the coming century by different modeling assumptions. We compared projections based on occupancy models against colonization–extinction models, conducting the modeling at alternative spatial scales and using fine‐ or coarse‐resolution deadwood data. We also tested effects of key explanatory variables on species occurrence and colonization–extinction dynamics. The hierarchical Bayesian models applied were fitted to an extensive repeated survey of deadwood and fungi at 174 patches. We projected higher occurrence probabilities and more positive trends using the occupancy models compared to the colonization–extinction models, with greater difference for the species with lower occupancy, colonization rate, and colonization:extinction ratio than for the species with higher estimates of these statistics. The magnitude of future increase in occupancy depended strongly on the spatial modeling scale and resource resolution. We encourage using colonization–extinction models over occupancy models, modeling the process at the finest resource‐unit resolution that is utilizable by the species, and conducting projections for the same spatial scale and resource resolution at which the model fitting is conducted. Further, the models applied should include key variables driving the metapopulation dynamics, such as the availability of suitable resource units, habitat quality, and spatial connectivity.     

Distribution of Mycobacterium avium subspecies paratuberculosis in the lower Florida Keys

Johne's disease, a fatal and contagious gastrointestinal infection caused by Mycobacterium avium subsp. paratuberculosis (Map), was first diagnosed in an endangered Florida Key deer (Odocoileus virginianus clavium) in 1996 and later in six additional Key deer deaths from 1998 to 2004. We investigated the geographic distribution of Map in the Lower Florida Keys from February 2005 through May 2006 via collection of blood and fecal pellets from 51 live-captured deer, collection of 550 fecal samples from the ground, and by necropsies of 90 carcasses. Tissue and fecal samples also were submitted from 30 raccoons (Procyon lotor), three feral cats (Felis catus), an opossum (Didelphis virginiana), and a Lower Keys marsh rabbit (Sylvilagus palustris hefneri). Mycobacterium avium subsp. paratuberculosis was identified in 23 Key deer fecal samples collected from the ground, tissue samples from two clinically ill Key deer, and from the mesenteric lymph node of a raccoon. The results of this study indicate that Map persists in the Key deer population and environment at a low prevalence, but its distribution currently is limited to a relatively small geographic area within the range of Key deer.     

Population estimation and monitoring of an endangered lagomorph

We conducted the most intensive estimate of the endangered Lower Keys marsh rabbit (Sylvilagus palustris hefneri) metapopulation to date using pellet surveys and capture–recapture methodology. We livetrapped 83 rabbits, evaluated 5 closed population models, and selected the model that best represented the data. We considered the variation in behavioral response model the best model and correlated (r² = 0.913) its patch population estimates to patch pellet densities. From the prediction equation, we generated a range-wide metapopulation estimate of 317 rabbits, a western clade population of 257 rabbits, an eastern clade population of 25 rabbits, and translocated marsh rabbit populations of 35 and zero on Little Pine and Water keys, respectively. A subset of patches whose marsh rabbit subpopulations were last estimated in 1993 exhibited a 46% decline in abundance over 15 yr. Due to the low estimate of the eastern clade population, special effort should be initiated to avoid loss of genetic diversity. The prediction equation suffers from limited data at high pellet densities, patches with ≥5 pellets/m². Future studies should investigate if the slope of the regression is indeed near 1 by sampling patches across the range of pellet densities, especially those with ≥5 pellets/m². The equation provides managers a quick, efficient, and noninvasive method to estimate marsh rabbit abundance from pellet counts but the confidence of predicted rabbit densities from high pellet density patches is low. © 2011 The Wildlife Society     

Invasive pythons,not anthropogenic stressors,explain the distribution of a keystone species

Untangling the causes of native species loss in human-modified systems is difficult and often controversial. Evaluating the impact of non-native species in these systems is particularly challenging, as additional human perturbations often precede or accompany introductions. One example is the ongoing debate over whether mammal declines within Everglades National Park (ENP) were caused by either the establishment of non-native Burmese pythons (Python molurus bivittatus) or the effects of other anthropogenic stressors. We examined the influence of both pythons and a host of alternative stressors—altered hydrology and habitat characteristics, mercury contamination and development—on the distribution of the marsh rabbit (Sylvilagus palustris), a once common mammal in ENP. Distance from the epicenter of the python invasion best explained marsh rabbit occurrence in suitable habitat patches, whereas none of the alternative stressors considered could explain marsh rabbit distribution. Estimates of the probability of marsh rabbit occurrence ranged from 0 at the python invasion epicenter to nearly 1.0 150 km from the invasion epicenter. These results support the hypothesis that invasive pythons shape the distribution of marsh rabbits in southern Florida. The loss of marsh rabbits and similar species will likely alter trophic interactions and ecosystem function within the Everglades, an internationally important hotspot of biodiversity. Further, our results suggest that non-native species can have profound impacts on mainland biodiversity.