Biotic invasions and habitat modification are two drivers of global change predicted to have detrimental impacts on the persistence of indigenous biota worldwide. Few studies have investigated how they operate synergistically to alter trophic interactions among indigenous and nonindigenous species in invaded ecosystems. We experimentally manipulated a suite of interacting invasive mammals, including top predators (cat Felis catus, ferret Mustela furo, stoat M. erminea), herbivores (rabbit Oryctolagus cuniculus, hare Lepus europaeus), and an insectivore (hedgehog Erinaceus europaeus occidentalis), and measured their effects on indigenous lizards and invertebrates and on an invasive mesopredator (house mouse Mus musculus). The work was carried out in a grassland/shrubland ecosystem that had been subjected to two types of habitat modification (widespread introduction of high-seed-producing pasture species, and areas of land use intensification by fertilization and livestock grazing). We also quantified food productivity for indigenous and invasive fauna by measuring pasture biomass, as well as seed and fruit production by grasses and shrubs. Indigenous fauna did not always increase following top-predator suppression: lizards increased on one of two sites; invertebrates did not increase on either site. Mesopredator release of mice was evident at the site where lizards did not increase, suggesting negative effects of mice on lizard populations. High mouse abundance occurred only on the predator-suppression site with regular production of pasture seed, indicating that this food resource was the main driver of mouse populations. Removal of herbivores increased pasture and seed production, which further enhanced ecological release of mice, particularly where pasture swards were overtopped by shrubs. An effect of landscape supplementation was also evident where nearby fertilized pastures boosted rabbit numbers and the associated top predators. Other studies have shown that both suppression of invasive predators and retiring land from grazing can benefit indigenous species, but our results suggest that the ensuing vegetation changes and complex interactions among invasive species can block recovery of indigenous fauna vulnerable to mesopredators. Top-down and bottom-up ecological release of mesopredators and landscape supplementation of top predators are key processes to consider when managing invaded communities in complex landscapes.

Summary1. Plant invasions are predicted to accelerate in a world with increased anthropogenic disturbance. Non-native species pre-adapted to these disturbances may especially be poised to invade novel communities. Conservation managers therefore need predictions of how to alter disturbances to maximize the persistence of native biodiversity. 2. We tested a multivariate hypothesis about the causal mechanisms underlying plant invasions in an ephemeral wetland in South Island, New Zealand, to inform management of this biodiverse but globally imperilled habitat. Our approach details among the first applications in ecology of Bayesian structural equation modelling, demonstrating its potential to inform management by disentangling the relative importance of strongly intercorrelated processes. 3. We found that invasion by non-native plants was lowest in sites where the physical disturbance caused by flooding was both intense and frequent. This effect was stronger than the positive response of non-native species to high soil N supply, which was positively related to flooding. 4. Sites flooded over a 4-year period had greater reductions in invasion than those associated with floods in the year prior to plot measurement because non-native species lacked traits for long-term persistence beneath water. Grazer exclusion had a small positive effect on invasion, as non-native species were preferentially selected by the herbivores at our site. 5. Our results show that only species adapted to the dominant disturbance regimes at a site may become successful invaders. Species native to ephemeral wetlands have specially evolved traits that allow them to persist and dominate in these sites. 6. Synthesis and applications. Predictions of invasions in a world of multiple disturbances clearly need to consider whether the evolutionary history of non-native species predisposes them to invade novel communities. Maintaining hydrological and nutrient regimes of ephemeral wetlands will limit the number of introduced species that are pre-adapted to become invasive.

Introduced mammalian predators threaten populations of endemic New Zealand skinks. Their effects on skink populations have been not often quantified on the mainland and are known primarily from skink population increases on islands from which mammals have been eradicated. Estimating skink population density with capture-recapture trapping is time-consuming and costly. Counting skinks in artificial retreats in specific weather conditions may be a useful and relatively quick way to index population density, but needs calibration for different habitats and species. In 2007 and 2009, we estimated the population density of small terrestrial skinks (McCann's skink Oligosoma maccanni, southern grass skink O. aff. polychroma clade 5 and cryptic skink O. inconspicuum), based on spatially explicit capture-recapture (SECR) in pitfall traps in three mammal-management treatments at Macraes Flat, Otago. The treatments were eradication of large predators and near-eradication of rodents inside a mammal-resistant fence, suppression of mammalian predator populations through continuous trapping (two locations within an extensive area), and no mammal management. We tested for relationships between the estimated population densities and dawn and late-morning counts of skinks in artificial retreats. Skink density (three species combined) ranged from c. 1200 per ha at the experimental control site to c. 4000 per ha at the fenced site. These treatment differences in skink density may be the combined effect of predator management and pre-existing differences due to habitat characteristics and farming practices. Skink counts done in late morning (2009) were related to estimated skink densities but did not differ significantly between treatments. Skink counts done at dawn (both years) were not related to densities. Counts, but not densities, were significantly higher at locations with a more northerly aspect. We recommend further investigation of the utility of skink counts in artificial retreats for monitoring skink density at this location, with careful control of ambient temperature during sampling, and of aspect, habitat and device placement.

House mice are among the most widely distributed mammals in the world, and adversely affect a wide range of indigenous biota. Suppressing mouse populations, however, is difficult and expensive. Cost-effective suppression requires knowing how low to reduce mouse numbers to achieve biodiversity outcomes, but these targets are usually unknown or not based on evidence. We derived density-impact functions (DIFs) for mice and small indigenous fauna in a tussock grass/shrubland ecosystem. We related two indices of mouse abundance to five indices of indigenous lizard and invertebrate abundance measured inside and outside mammal-resistant fences. Eight of 22 DIFs were significantly non-linear, with positive responses of skinks (Oligosoma maccanni, O. polychroma) and ground wētā (Hemiandrus spp.) only where mice were not detected or scarce (< 5% footprint tunnel tracking rate or printing rate based on footprint density). Kōrero geckos (Woodworthia spp.) were rarely detected where mice were present. A further 9 DIFs were not differentiated from null models, but patterns were consistent with impacts at 5% mouse abundance. This study suggests that unless mouse control programmes commit to very low abundances, they risk little return for effort. Impact studies of invasive house mice are largely restricted to island ecosystems. Studies need to be extended to other ecosystems and species to confirm the universality or otherwise of these highly non-linear DIFs.

Abstract:Estimates of vegetation attributes measured by sampling often inform scientific inference, management actions, and policy decisions. However, different sampling methods and sample sizes (i.e. number of plots) can yield significantly different estimates of vegetation attributes. This occurs because the abundance distributions and spatial distributions of species in the plant community influence their probabilities of detection and estimates of their abundances. We predicted that different sampling methods and sample sizes would produce significantly different estimates not only of vascular plant species diversity, but also of indigenous dominance (the level of indigenous influence) in mixed vegetation where indigenous and exotic floras have different abundance or spatial distributions. To test our predictions we applied three sampling methods to 24 plots in grassland and cushion vegetation in a 1058-ha scientific reserve in the Upper Waitaki (Mackenzie) Basin, New Zealand. Our methods sampled ground areas from 0.65 to 400 m 2 , and included two variants of common 'subsampling' approaches, which assessed only discrete subunits within larger plots. Indigenous plant species were both less abundant on average and more spatially-clustered (i.e. less evenly dispersed across plots) than exotic species. The two subsampling methods were less likely to detect less abundant and more spatially clustered species, leading to lower ratios of indigenous to exotic species recorded, and lower estimates of indigenous dominance of composition (% of species indigenous). Numbers of indigenous species accumulated more rapidly with increasing sample size than numbers of exotic species, so that indigenous dominance also increased with the number of plots sampled. We conclude that properly measuring species diversity and indigenous dominance in mixed indigenous-exotic plant communities requires both the searching of sizeable plots and use of rarefaction rather than plot-averaging of statistics. We suggest greater use of rarefaction and more consideration of species' detection probabilities in sampling New Zealand's mixed indigenous-exotic plant communities should improve the reliability, transparency and comparability of measures of diversity and may also provide new ecological insights.

Factors controlling vegetation restoration of depleted short-tussock grasslands are poorly understood. We investigated effects of mouse-ear hawkweed ('hawkweed', Pilosella officinarum) cover and environmental stress associated with landform and soil type on the rate and pattern of indigenous vegetation recovery from grazing in the highly-modified 1000-ha Lake Tekapo Scientific Reserve in the north of the Upper Waitaki ('Mackenzie') Basin. The reserve has been destocked of sheep and under effective rabbit control since 1992. At that time, mouse-ear hawkweed dominated vegetation on three of its five major landforms (up to 42% cover), and 44-89% of soil was exposed. In 2011 we resampled 12 original vegetation monitoring plots that were established in 1993. Indigenous vegetation recovered in the 18 years following removal from grazing despite high levels of initial modification and exotic cover dominance. Exposed soil and rock decreased, and indigenous plant cover and litter increased across all landforms, while vascular and nonvascular indigenous plant cover increased at different ends of a landform-driven productivity gradient. Hawkweed invasion did not retard recovery; to the contrary, the extent and rate of recovery was higher on more productive landforms with higher initial hawkweed cover. The pattern of change across the reserve was consistent with grazing having exerted a powerful constraint on the growth and biomass of both indigenous and exotic palatable species prior to reservation. Soil moisture stress appeared to delay the timing of hawkweed invasion, and to constrain productivity and hence the rate of indigenous vegetation recovery following release from grazing. We propose a testable model of interacting influences of grazing and environment on indigenous vegetation and the niche of mouse-ear hawkweed, in which recovery outcomes depend on environmental productivity. We suggest that removal of feral grazing as well as stock, size of recovery area, limited monitoring-disturbance, and timing of grazing-release relative to hawkweed invasion may explain why our results and conclusions contrast with those from other studies of release of fescue-tussock grasslands from pastoral grazing.