Invasive-dominated grasslands in Hawai'i are resilient to disturbance.

Non-native-dominated landscapes may arise from invasion by competitive plant species, disturbance and invasion of early-colonizing species, or some combination of these. Without knowing site history, however, it is difficult to predict how native or non-native communities will reassemble after disturbance events. Given increasing disturbance levels across anthropogenically impacted landscapes, predictive understanding of these patterns is important. We asked how disturbance affected community assembly in six invaded habitat types common in dryland, grazed landscapes on Island of Hawai'i. We mechanically disturbed 100 m2 plots in six vegetation types dominated by one of four invasive perennial grasses (Cenchrus ciliaris, Cenchrus clandestinus, Cenchrus setaceus, or Melinis repens), a native shrub (Dodonaea viscosa), or a native perennial bunchgrass (Eragrostis atropioides). We censused vegetation before disturbance and monitored woody plant colonization and herbaceous cover for 21 months following the disturbance, categorizing species as competitors, colonizers, or a combination, based on recovery patterns. In addition, we planted individuals of the native shrub and bunchgrass and monitored survival to overcome dispersal limitation of native species when exploring these patterns. We found that the dominant vegetation types showed variation in post-disturbance syndrome, and that the variation in colonizer versus competitor syndrome occurred both between species, but also within species among different vegetation types. Although there were flushes of native shrub seedlings, these did not survive to 21 months within invaded habitats, probably due to regrowth by competitive invasive grasses. Similarly, survival of planted native individuals was related to the rate of regrowth by dominant species. Regardless of colonization/competitor syndrome, however, all dominant vegetation types were relatively resilient to change. Our results highlight that the altered post-agricultural, invaded grassland landscapes in Hawai'i are stable states. More generally, they point to the importance of resident communities and their effects on species interactions and seed availability in shaping plant community response to disturbance.

Crossing the threshold: Invasive grasses inhibit forest restoration on Hawaiian islands.

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 Hawai'i 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 ('ohi'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 'ohi'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 'ohi'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.

Fire modifies plant-soil feedbacks.

Although plant-soil feedbacks (interactions between plants and soils, often mediated by soil microbes, abbreviated as PSFs) are widely known to influence patterns of plant diversity at local and landscape scales, these interactions are rarely examined in the context of important environmental factors. Resolving the roles of environmental factors is important because the environmental context may alter PSF patterns by modifying the strength or even direction of PSFs for certain species. One important environmental factor that is increasing in scale and frequency with climate change is fire, though the influence of fire on PSFs remains essentially unexamined. By changing microbial community composition, fire may alter the microbes available to colonize the roots of plants and thus seedling growth post-fire. This has potential to change the strength and/or direction of PSFs, depending on how such changes in microbial community composition occur and the plant species with which the microbes interact. We examined how a recent fire altered PSFs of two leguminous, nitrogen-fixing tree species in Hawai'i. For both species, growing in conspecific soil resulted in higher plant performance (as measured by biomass production) than growing in heterospecific soil. This pattern was mediated by nodule formation, an important process for growth for legume species. Fire weakened PSFs for these species and therefore pairwise PSFs, which were significant in unburned soils, but were nonsignificant in burned soils. Theory suggests that positive PSFs such as those found in unburned sites would reinforce the dominance of species where they are locally dominant. The change in pairwise PSFs with burn status shows PSF-mediated dominance might diminish after fire. Our results demonstrate that fire can modify PSFs by weakening the legume-rhizobia symbiosis, which may alter local competitive dynamics between two canopy dominant tree species. These findings illustrate the importance of considering environmental context when evaluating the role of PSFs for plants.

The role of microtopography and resident species in post-disturbance recovery of arid habitats in Hawai'i.

Habitat-suitability indices (HSI) have been employed in restoration to identify optimal sites for planting native species. Often, HSI are based on abiotic variables and do not include biotic interactions, even though similar abiotic conditions can favor both native and nonnative species. Biotic interactions such as competition may be especially important in invader-dominated habitats because invasive species often have fast growth rates and can exploit resources quickly. In this study, we test the utility of an HSI of microtopography derived from airborne LiDAR to predict post-disturbance recovery and native planting success in native shrub-dominated and nonnative, invasive grass-dominated dryland habitats in Hawai'i. The HSI uses high-resolution digital terrain models to classify sites' microtopography as high, medium, or low suitability, based on wind exposure and topographic position. We used a split-plot before-after-control-impact design to implement a disturbance experiment within native shrub (Dodonaea viscosa) and nonnative, invasive grass (Cenchrus clandestinus)-dominated ecosystems across three microtopography categories. In contrast to previous studies using the same HSI, we found that microtopography was a poor predictor of pre-disturbance conditions for soil nutrients, organic matter content, or foliar C:N, within both Dodonaea and Cenchrus vegetation types. In invader-dominated Cenchrus plots, microtopography helped predict cover, but not as expected (i.e., highest cover would be in high-suitability plots): D. viscosa had the greatest cover in low-suitability and C. clandestinus had the greatest cover in medium-suitability plots. Similarly, in native-dominated Dodonaea plots, microtopography was a poor predictor of D. viscosa, C. clandestinus, and total plant cover. Although we found some evidence that microtopography helped inform post-disturbance plant recovery of D. viscosa and total plant cover, vegetation type was a more important predictor. Important for considering the success of plantings, percent cover of D. viscosa decreased while percent cover of C. clandestinus increased within both vegetation types 20 months after disturbance. Our results are evidence that HSIs based on topographic features may prove most useful for choosing planting sites in harsh habitats or those already dominated by native species. In more productive habitats, competition from resident species may offset any benefits gained from "better" suitability sites.

Can the impact of canopy trees on soil and understory be altered using litter additions?

Trees can have large effects on soil nutrients in ways that alter succession, particularly in the case of nitrogen-(N)-fixing trees. In Hawai'i, 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 ('ohi'a), the dominant native tree in Hawai'i, 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 'ohi'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 'ohi'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 'ohi'a litter increased soil carbon, but increased rather than decreased inorganic soil N pools. Contrary to expectations, koa litter decomposed more slowly than 'ohi'a, although it released more N per unit of litter. We saw no reduction in grass biomass due to 'ohi'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.

Environmental gradients influence differences in leaf functional traits between native and non-native plants.

Determining the characteristics of non-native plants that can successfully establish and spread is central to pressing questions in invasion ecology. Evidence suggests that some non-native species establish and spread in new environments because they possess characteristics (functional traits) that allow them to either successfully compete with native residents or fill previously unfilled niches. However, the relative importance of out-competing native species vs. filling empty niche space as potential mechanisms of invasion may depend on environmental characteristics. Here, we measured plant functional traits, proxies indicative of competitive and establishment strategies, to determine if these traits vary among native and invasive species and if their prevalence is dependent on environmental conditions. Using a natural environmental gradient in Hawai'i Volcanoes National Park, we evaluated how functional traits differ between native and non-native plant communities and if these differences change along an environmental gradient from hot, dry to cool, wet conditions. Functional trait differences suggested that both competition and open niche space may be important for invasion. Non-native communities tended to have traits associated with faster growth strategies such as higher specific leaf area and lower leaf thickness. However, native and non-native community traits became more dissimilar along the gradient, suggesting that non-native species may be occupying previously unfilled niche space at the hot, dry end of the gradient. We also found that most of the variation in functional trait values amongst plots was due to species turnover rather than intraspecific variation. These results highlight the role of environmental context when considering invasion mechanisms.

Invasive rat control is an efficient, yet insufficient, method for recovery of the critically endangered Hawaiian plant hau kuahiwi (Hibiscadelphus giffardianus).

Biological invasions of rodents and other species have been especially problematic on tropical islands. Invasive Rattus rattus consumption of Hibiscadelphus giffardianus (Malvaceae; common Hawaiian name hau kuahiwi) fruit and seeds has been hypothesized to be the most-limiting factor inhibiting the critically endangered tree, but this has not been experimentally tested, and little is known about other factors affecting seed dispersal, germination, and seedling establishment. Thus, we do not know if rat removal is sufficient to increase hau kuahiwi recruitment. This study aims to evaluate the effect of rat population control on the ability of hau kuahiwi to retain fruit and establish seedlings. We compared hau kuahiwi fruiting and seedling recruitment in a stand treated to reduce rat abundance and a neighbouring control stand. Fruit retention increased following treatment but seedling establishment did not. Although rat control improves the ability of hau kuahiwi to retain fruit, other, presently unknown inhibitors to seed dispersal, germination, and/or seedling development remain. Seed and seedling predation by other species, competition from numerous invasive plant species, unsuitable climate, and/or other factors may be primary inhibitors in the absence of rats, but we emphasize that progressive isolation of these factors at individual hau kuahiwi life stages may be necessary to identify the remaining threats to the conservation of this critically endangered plant.

Evaluating nurse plants for restoring native woody species to degraded subtropical woodlands.

Harsh habitats dominated by invasive species are difficult to restore. Invasive grasses in arid environments slow succession toward more desired composition, yet grass removal exacerbates high light and temperature, making the use of "nurse plants" an appealing strategy. In this study of degraded subtropical woodlands dominated by alien grasses in Hawai'i, we evaluated whether individuals of two native (Dodonaea viscosa, Leptocophylla tameiameia) and one non-native (Morella faya) woody species (1) act as natural nodes of recruitment for native woody species and (2) can be used to enhance survivorship of outplanted native woody species. To address these questions, we quantified the presence and persistence of seedlings naturally recruiting beneath adult nurse shrubs and compared survival and growth of experimentally outplanted seedlings of seven native woody species under the nurse species compared to intact and cleared alien-grass plots. We found that the two native nurse shrubs recruit their own offspring, but do not act as establishment nodes for other species. Morella faya recruited even fewer seedlings than native shrubs. Thus, outplanting will be necessary to increase abundance and diversity of native woody species. Outplant survival was the highest under shrubs compared to away from them with few differences between nurse species. The worst habitat for native seedling survival and growth was within the unmanaged invasive grass matrix. Although the two native nurse species did not differentially affect outplant survival, D. viscosa is the most widespread and easily propagated and is thus more likely to be useful as an initial nurse species. The outplanted species showed variable responses to nurse habitats that we attribute to resource requirements resulting from their typical successional stage and nitrogen fixation capability.

A mechanistic study of plant and microbial controls over R* for nitrogen in an annual grassland.

Differences in species' abilities to capture resources can drive competitive hierarchies, successional dynamics, community diversity, and invasions. To investigate mechanisms of resource competition within a nitrogen (N) limited California grassland community, we established a manipulative experiment using an R* framework. R* theory holds that better competitors within a N limited community should better depress available N in monoculture plots and obtain higher abundance in mixture plots. We asked whether (1) plant uptake or (2) plant species influences on microbial dynamics were the primary drivers of available soil N levels in this system where N structures plant communities. To disentangle the relative roles of plant uptake and microbially-mediated processes in resource competition, we quantified soil N dynamics as well as N pools in plant and microbial biomass in monoculture plots of 11 native or exotic annual grassland plants over one growing season. We found a negative correlation between plant N content and soil dissolved inorganic nitrogen (DIN, our measure of R*), suggesting that plant uptake drives R*. In contrast, we found no relationship between microbial biomass N or potential net N mineralization and DIN. We conclude that while plant-microbial interactions may have altered the overall quantity of N that plants take up, the relationship between species' abundance and available N in monoculture was largely driven by plant N uptake in this first year of growth.

Self-reinforcing impacts of plant invasions change over time.

Returning native species to habitats degraded by biological invasions is a critical conservation goal. A leading hypothesis poses that exotic plant dominance is self-reinforced by impacts on ecosystem processes, leading to persistent stable states. 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. However, few studies have followed invasions through time to ask whether ecosystem impacts and feedbacks persist. Here we return to woodland sites in Hawai'i Volcanoes National Park that were invaded by exotic C4 grasses in the 1960s, the ecosystem impacts of which were studied intensively in the 1990s. 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. 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.

Regional constraints to biological nitrogen fixation in post-fire forest communities.

Biological nitrogen fixation (BNF) is a key ecological process that can restore nitrogen (N) lost in wildfire and shape the pace and pattern of post-fire forest recovery. To date, there is limited information on how climate and soil fertility interact to influence different pathways of BNF in early forest succession. We studied asymbiotic (forest floor and soil) and symbiotic (the shrub Ceanothus integerrimus) BNF rates across six sites in the Klamath National Forest, California, USA. We used combined gradient and experimental phosphorus (P) fertilization studies to explore cross-site variation in BNF rates and then related these rates to abiotic and biotic variables. We estimate that our measured BNF rates 22 years after wildfire (6.1-12.1 kg N x ha(-1) x yr(-1)) are unlikely to fully replace wildfire N losses. We found that asymbiotic BNF is P limited, although this is not the case for symbiotic BNF in Ceanothus. In contrast, Ceanothus BNF is largely driven by competition from other vegetation: in high-productivity sites with high potential evapotranspiration (Et), shrub biomass is suppressed as tree biomass increases. Because shrub biomass governed cross-site variation in Ceanothus BNF, this competitive interaction led to lower BNF in sites with high productivity and Et. Overall, these results suggest that the effects of nutrients play a larger role in driving asymbiotic than symbiotic fixation across our post-fire sites. However, because symbiotic BNF is 8-90x greater than asymbiotic BNF, it is interspecific plant competition that governs overall BNF inputs in these forests.

The role of plant-soil feedbacks in driving native-species recovery.

The impacts of exotic plants on soil nutrient cycling are often hypothesized to reinforce their dominance, but this mechanism is rarely tested, especially in relation to other ecological factors. In this manuscript we evaluate the influence of biogeochemically mediated plant-soil feedbacks on native shrub recovery in an invaded island ecosystem. The introduction of exotic grasses and grazing to Santa Cruz Island, California, USA, converted native shrublands (dominated by Artemisia californica and Eriogonum arborescens) into exotic-dominated grasslands (dominated by Avena barbata) over a century ago, altering nutrient-cycling regimes. To test the hypothesis that exotic grass impacts on soils alter reestablishment of native plants, we implemented a field-based soil transplant experiment in three years that varied widely in rainfall. Our results showed that growth of Avena and Artemisia seedlings was greater on soils influenced by their heterospecific competitor. Theory suggests that the resulting plant-soil feedback should facilitate the recovery of Artemisia in grasslands, although four years of monitoring showed no such recovery, despite ample seed rain. By contrast, we found that species effects on soils lead to weak to negligible feedbacks for Eriogonum arborescens, yet this shrub readily colonized the grasslands. Thus, plant-soil feedbacks quantified under natural climate and competitive conditions did not match native-plant recovery patterns. We also found that feedbacks changed with climate and competition regimes, and that these latter factors generally had stronger effects on seedling growth than species effects on soils. We conclude that even when plant-soil feedbacks influence the balance between native and exotic species, their influence may be small relative to other ecological processes.

California annual grass invaders: the drivers or passengers of change?

The dominance of invasive species is often assumed to reflect their competitive superiority over displaced native species. However, invasive species may be abundant because of their greater tolerance to anthropogenic impacts accompanying their introduction. Thus, invasive species can either be the drivers or passengers of change.We distinguish between these two possibilities in California grasslands currently dominated by Mediterranean annuals (exotics) and subjected to livestock grazing since European settlement. We focused on native annual grasses and forbs, an understudied species-rich component of the California flora, and Mediterranean annual grasses, currently dominant and among the first non-native plants introduced to the area.We established a field experiment with fenced and unfenced blocks in a cattle pasture. We measured concentrations of limiting resources (nitrogen, phosphorus, light and soil moisture) in monoculture plots as an index of competitive ability (i.e. R*). We then quantified grazing impacts on biomass and seed production in grazed vs. ungrazed monoculture plots. Finally, we measured biomass and seed production of each species competing in mixture plots, in the presence and absence of grazers.We found that native and exotic species did not differ in R* indices of competitive ability, i.e. concentrations of limiting resources in ungrazed native monoculture plots did not differ from concentrations in ungrazed exotic monoculture plots. By contrast, exotic annuals suffered less from grazing than native annuals, perhaps reflecting their longer evolutionary history with cattle grazing. Consistent with these results, native and exotic annuals were equally abundant in ungrazed mixtures, but exotic species overwhelmingly dominated grazed mixtures.Species able to draw down nitrogen and light to lower levels in monocultures (i.e. those with lower R* values) dominated biomass and seeds in mixed plots without grazers. However, R* did not predict the relative abundance of species in grazed plots. Moreover, the relative abundance of species in mixtures did not correlate with grazing impacts on their monocultures, implying that grazing alters inter-specific competitive dynamics.Synthesis. We demonstrate that the displacement of native annuals by Mediterranean annual grasses in California may largely have been driven by cattle grazing.

Plant-soil feedbacks and invasive spread.

Plant invaders have been suggested to change soil microbial communities and biogeochemical cycling in ways that can feedback to benefit themselves. In this paper, we ask when do these feedbacks influence the spread of exotic plants. Because answering this question is empirically challenging, we show how ecological theory on 'pushed' and 'pulled' invasions can be used to examine the problem. We incorporate soil feedbacks into annual plant invasion models, derive the conditions under which such feedbacks affect spread, and support our approach with simulations. We show that in homogeneous landscapes, strong positive feedbacks can influence spreading velocity for annual invaders, but that empirically documented feedbacks are not strong enough to do so. Moreover, to influence spread, invaders must modify the soil environment over a spatial scale larger than is biologically realistic. Though unimportant for annual invader spread in our models, feedbacks do affect invader density and potential impact. We discuss how future research might consider the way landscape structure, dispersal patterns, and the time scales over which plant-soil feedbacks develop regulate the effects of such feedbacks on invader spread.