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Plant invasions are thought to alter the ecosystem in a way that disadvantages the native species, making re-establishment after eradication difficult; here, on returning to a site at which an invasive plant altered nitrogen-mineralization levels several decades ago, mineralization is found to have returned to pre-invasion levels, although these new conditions favour new invaders over the natives. New invaders for old Invasive plants can radically change ecosystem function to the detriment of species originally in place. But what happens if the invasive species is eradicated — is the ecosystem changed permanently, or if the ousted plants are replaced can they carry on as before? For this study Stephanie Yelenik and Carla D'Antonio returned to grass-invaded field sites in Hawai'i Volcanoes National Park to study invader impacts replicated more than two decades on. The invasive grass had altered nitrogen-mineralization levels but decades later the ecosystem had returned to pre-invasion levels. However, the new conditions did not favour native re-establishment; rather, they benefited a new invasive, the nitrogen-fixing tree Morella faya . Returning native species to habitats degraded by biological invasions is a critical conservation goal 1 . A leading hypothesis poses that exotic plant dominance is self-reinforced by impacts on ecosystem processes, leading to persistent stable states 2 , 3 , 4 , 5 , 6 . Invaders have been documented to modify fire regimes, alter soil nutrients or shift microbial communities in ways that feed back to benefit themselves over competitors 2 , 5 , 6 , 7 . However, few studies have followed invasions through time to ask whether ecosystem impacts and feedbacks persist 8 , 9 . Here we return to woodland sites in Hawai′i Volcanoes National Park that were invaded by exotic C 4 grasses in the 1960s, the ecosystem impacts of which were studied intensively in the 1990s 10 , 11 , 12 . We show that positive feedbacks between exotic grasses and soil nitrogen cycling have broken down, but rather than facilitating native vegetation, the weakening feedbacks facilitate new exotic species. Data from the 1990s showed that exotic grasses increased nitrogen-mineralization rates by two- to fourfold, but were nitrogen-limited 10 , 12 , 13 . Thus, the impacts of the invader created a positive feedback early in the invasion. We now show that annual net soil nitrogen mineralization has since dropped to pre-invasion levels. In addition, a seedling outplanting experiment that varied soil nitrogen and grass competition demonstrates that the changing impacts of grasses do not favour native species re-establishment. Instead, decreased nitrogen availability most benefits another aggressive invader, the nitrogen-fixing tree Morella faya . Long-term studies of invasions may reveal that ecosystem impacts and feedbacks shift over time, but that this may not benefit native species recovery.

While the influence of canopy trees on soils in natural and restored forest environments is well studied, the influence of understory species is not. Here, we evaluate the effects of outplanted native woody understory on invasive grass biomass and soil nutrient properties in heavily grass-invaded 30 + year-old plantations of a native N-fixing tree Acacia koa in Hawai‘i. We analyze soils from under A. koa trees with versus without planted woody understory and compare these to soils from under remnant pasture trees of the pre-deforestation dominant, Metrosideros polymorpha where passive recruitment of native woody understory has occurred since the cessation of grazing. Simultaneously, we experimentally planted understory species at three times the density used by managers to see if this could quickly decrease grass biomass and change soil nutrient dynamics. We found that invasive grass biomass declined with understory planting in surveyed and experimental sites. Yet, woody understory abundance had no effect on N cycling. Short-term N availability and nitrification potential were higher under A. koa than M. polymorpha trees regardless of understory. Net N mineralization either did not differ (~ 1 mo) between canopy species or was higher (171 day incubations) under remnant M. polymorpha where organic matter was also higher. The only influence of understory on soil was a positive correlation with loss-on-ignition (organic matter) under M. polymorpha. We also demonstrate differential controls over N cycling under the two canopy tree species. Overall, understory restoration has not changed soil characteristics even as invasive grass biomass declines.

Humans and their ancestors are unique in being a fire-making species, but 'natural' (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human-set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from ' natural' background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke-related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas-induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1 ) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre-human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.

Invasion by exotic plant species can profoundly affect native plant species performance and the inferred proximate cause is typically competition. We used invaded grasslands in the semi-arid Western USA to separate resource competition from structural interference of an exotic grass on native forb performance, specifically evaluating the role of competition from living vegetation versus litter accumulation (hereafter, thatch). We simultaneously tested whether a positive thatch-feedback exists for the dominant exotic grass species, Bromus diandrus. Thatch and B. diandrus density were manipulated and coupled with native seed addition to separate the effect of B. diandrus competition relative to thatch accumulation. To determine the response of native forb species and B. diandrus to varying thatch abundance, we created a gradient of thatch densities and measured species response, soil moisture and light availability. The thatch of B. diandrus greatly reduced native forb performance (number of individuals and biomass) and resulted in near complete exclusion of many species. The effect of living B. diandrus density on native forb performance was minimal and inconsistent. Forb performance and light availability both exhibited exponential declines with thatch build-up suggesting that light reduction is a primary mechanism through which thatch affects forbs. Simultaneous with forb suppression, B. diandrus performance was positively affected by thatch, consistent with a positive feedback initiated by invasion of this species. Our results demonstrate that thatch accumulation, rather than competition for resources, is the primary cause of native species decline in grasslands invaded by this exotic annual grass. In addition, the dominance of B. diandrus in many invaded grasslands may be reinforced through a positive thatch-feedback and require active management, such as grazing or fire, to be broken.

Increased fire frequency has been shown to promote alien plant invasions in the western United States, resulting in persistent vegetation type change. Short interval fires are widely considered to be detrimental to reestablishment of shrub species in southern California chaparral, facilitating the invasion of exotic annuals and producing \"type conversion\". However, supporting evidence for type conversion has largely been at local, site scales and over short post-fire time scales. Type conversion has not been shown to be persistent or widespread in chaparral, and past range improvement studies present evidence that chaparral type conversion may be difficult and a relatively rare phenomenon across the landscape. With the aid of remote sensing data covering coastal southern California and a historical wildfire dataset, the effects of short interval fires (<8 years) on chaparral recovery were evaluated by comparing areas that burned twice to adjacent areas burned only once. Twelve pairs of once- and twice-burned areas were compared using normalized burn ratio (NBR) distributions. Correlations between measures of recovery and explanatory factors (fire history, climate and elevation) were analyzed by linear regression. Reduced vegetation cover was found in some lower elevation areas that were burned twice in short interval fires, where non-sprouting species are more common. However, extensive type conversion of chaparral to grassland was not evident in this study. Most variables, with the exception of elevation, were moderately or poorly correlated with differences in vegetation recovery.

Forest removal for livestock grazing is a striking example of human-caused state change leading to a stable, undesirable invasive grass system that is resistant to restoration efforts. Understanding which factors lead to resilience to the alternative grass state can greatly benefit managers when planning forest restoration. We address how thresholds of grass cover and seed rain might influence forest recovery in a restoration project on Hawaiʻi Island, USA. Since the 1980s, over 400,000 Acacia koa (koa) trees have been planted across degraded pasture, and invasive grasses still dominate the understory with no native woody-plant recruitment. Between this koa/grass matrix are remnant native Metrosideros polymorpha (‘ōhi‘a) trees beneath which native woody plants naturally recruit. We tested whether there were threshold levels of native woody understory that accelerate recruitment under both tree species by monitoring seed rain at 40 trees (20 koa and ‘ōhi‘a) with a range of native woody understory basal area (BA). We found a positive relationship between total seed rain (but not bird-dispersed seed rain) and native woody BA and a negative relationship between native woody BA and grass cover, with no indication of threshold dynamics. We also experimentally combined grass removal levels with seed rain density (six levels) of two common understory species in plots under koa (n = 9) and remnant ‘ōhi‘a (n = 9). Few seedlings emerged when no grass was removed despite adding seeds at densities two to 75 times higher than naturally occurring. However, seedling recruitment increased two to three times once at least 50% of grass was removed. Existing survey data of naturally occurring seedlings also supported a threshold of grass cover below which seedlings were able to establish. Thus, removal of all grasses is not necessary to achieve system responses: Even moderate reductions (~50%) can increase rates of native woody recruitment. The nonlinear thresholds found here highlight how incremental changes to an inhibitory factor lead to limited restoration success until a threshold is crossed. The resources needed to fully eradicate an invasive species may be unwarranted for state change, making understanding where thresholds lie of the utmost importance to prioritize resources.

Exotic annual grasses can dominate ecosystems by producing a thick layer of dead plant litter, hereafter “thatch,” which promotes the regeneration of exotic grasses and inhibits native plants. Vernal pool wetlands within a grassland matrix are threatened by these exotic annual grasses, meriting the need for long‐term management. We utilized the investment of local community members to test the efficacy of long‐term thatch management on urban vernal pool plant assemblages. We recruited over 40 undergraduate students to perform manual annual summer thatch removal around the edges of 15 urban vernal pools for four years. We coupled thatch removal with annual native seed addition because our analysis of environmental DNA (“eDNA”) in the soil seed bank revealed a lack of native plant species and an abundance of exotic plant species. Our annual thatch removal treatment successfully reduced thatch accumulation and increased bare ground, but it did not result in a consistent decrease in exotic plant cover or an increase in native plant cover. Instead, the effects of thatch manipulation on plant composition were modulated by annual precipitation, with exotic species increasing during dry years and native species increasing during wet years. The addition of native plant seed caused an increase in native plant species richness, but only after three years of annual thatch removal and native seed addition. Our results indicate that the restoration of native vernal pool plants can be limited by invasive species, native seed availability, and annual precipitation. Our findings show how engaging the local community in the long‐term restoration of urban ecosystems can address the persistent threat of invasion and build up capacity for native plant populations to increase over time.

The increased occurrences of drought and fire may be contributing to the loss of biodiverse ecosystems in Mediterranean regions. Specifically, the conversion of diverse native shrublands, such as chaparral, to non‐native annual grassland by fire is of great conservation concern in California. To avoid or slow the loss of chaparral, it is important to understand the underlying causes of landscape conversion. Studies investigating the interaction of multiple potential drivers are particularly crucial to identification of vulnerable areas of the landscape. Here we used aerial imagery to evaluate vegetation transitions between chaparral, sage scrub, grassland, and tree domination and their potential drivers within Ventura County, CA, a strongly Mediterranean climate region. We used random forest algorithms and conditional inference trees to determine the climatic, topographic, and fire‐related variables contributing most to vegetation change. Our results support that chaparral conversion to grass (27% of chaparral plots) is a result of landscape position, fire, and drought acting in tandem. In particular, lower elevation, southwest‐facing slopes that experience a postfire drought are at very high likelihood of conversion to non‐native annual grass. Additionally, our results show that these grasslands, once formed, rarely convert to other community types. Therefore, protecting shrub‐dominated areas that are most likely to convert (low elevation, more southwest‐facing slopes, and less annual precipitation) is crucial to preserving native vegetation diversity.

Trees can have large effects on soil nutrients in ways that alter succession, particularly in the case of nitrogen-(N)-fixing trees. In Hawaiʻi, forest restoration relies heavily on use of a native N-fixing tree, Acacia koa (koa), but this species increases soil-available N and likely facilitates competitive dominance of exotic pasture grasses. In contrast, Metrosideros polymorpha (‘ōhi‘a), the dominant native tree in Hawaiʻi, is less often planted because it is slow growing; yet it is typically associated with lower soil N and grass biomass, and greater native understory recruitment. We experimentally tested whether it is possible to reverse high soil N under koa by adding ‘ōhi‘a litter, using additions of koa litter or no litter as controls, over 2.5 yr. We then quantified natural litterfall and decomposition rates of ‘ōhi‘a and koa litter to place litter additions in perspective. Finally, we quantified whether litter additions altered grass biomass and if this had effects on native outplants. Adding ‘ōhi‘a litter increased soil carbon, but increased rather than decreased inorganic soil N pools. Contrary to expectations, koa litter decomposed more slowly than ‘ōhi‘a, although it released more N per unit of litter. We saw no reduction in grass biomass due to ‘ōhi‘a litter addition, and no change in native outplanted understory survival or growth. We conclude that the high N soil conditions under koa are difficult to reverse. However, we also found that outplanted native woody species were able to decrease exotic grass biomass over time, regardless of the litter environment, making this a better strategy for lowering exotic species impacts.

• Background and Aims Global environmental change will affect non-native plant invasions, with profound potential impacts on native plant populations, communities and ecosystems. In this context, we review plant functional traits, particularly those that drive invader abundance (invasiveness) and impacts, as well as the integration of these traits across multiple ecological scales, and as a basis for restoration and management. • Scope We review the concepts and terminology surrounding functional traits and how functional traits influence processes at the individual level. We explore how phenotypic plasticity may lead to rapid evolution of novel traits facilitating invasiveness in changing environments and then 'scale up' to evaluate the relative importance of demographic traits and their links to invasion rates. We then suggest a functional trait framework for assessing per capita effects and, ultimately, impacts of invasive plants on plant communities and ecosystems. Lastly, we focus on the role of functional trait-based approaches in invasive species management and restoration in the context of rapid, global environmental change. • Conclusions To understand how the abundance and impacts of invasive plants will respond to rapid environmental changes it is essential to link trait-based responses of invaders to changes in community and ecosystem properties. To do so requires a comprehensive effort that considers dynamic environmental controls and a targeted approach to understand key functional traits driving both invader abundance and impacts. If we are to predict ftiture invasions, manage those at hand and use restoration technology to mitigate invasive species impacts, future research must focus on functional traits that promote invasiveness and invader impacts under changing conditions, and integrate major factors driving invasions from individual to ecosystem levels.