Ecological Assessment of Dune Restorations in the Great Lakes Region

Because of the economic and environmental importance of stabilizing fragile sand dune habitats, restoration of dunes has become a common practice. Restoration efforts in the Great Lakes and East Coast regions of North America often consist of planting monocultures of the dominant native grass species, Ammophila breviligulata. We evaluated 18 dune restoration projects in the Great Lakes region conducted over the past 25 years. We characterized attributes of diversity (plants and insects), vegetation structure (plant biomass and cover), and ecological processes (soil nutrients and mycorrhizal fungi abundance) in each restoration, and we compared these measures to geographically paired natural dune communities. Restoration sites were similar to reference sites in most measured variables. Differences between restorations and reference sites were mostly explained by differences in ages, with the younger sites supporting slightly lower plant diversity and mycorrhizal spore abundance than older sites. Plant community composition varied little between restored and reference sites, with only one native forb species, Artemisia campestris, occurring significantly more often in reference sites than restored sites. Although it remains unclear whether more diverse restoration plantings could accelerate convergence on the ecological conditions of reference dunes, in general, traditional restoration efforts involving monoculture plantings of A. breviligulata in Great Lakes sand dunes appear to achieve ecological conditions found in reference dunes.     

Restoration of Nontarget Species: Bee Communities and Pollination Function in Riparian Forests

Nontarget species such as pollinators may be of great importance to the restoration process and the long‐term functioning of restored habitats, but little is known about how such groups respond to habitat restoration. I surveyed bee communities at five equal‐aged restored sites, paired with five reference sites (riparian remnants) along the Sacramento River, California, United States. Flower availability and bee visitation patterns were also measured to examine the restoration of pollination function. Restoration of structural vegetation allowed diverse and abundant native bee communities to establish at the restoration sites; however, the composition of these important pollinator communities was distinct from that in the remnant riparian sites. Differences did not arise primarily from differences in the composition of the flowering‐plant community; rather there must be other physical characteristics of the restored sites or differences in nesting site availability that led to the different pollinator communities. Because sites were spatially paired, the differences are unlikely to be driven by landscape context. Bee life‐history and other biological traits may partially explain the differences between bee communities at restored and remnant sites. Patterns of visitation to native plant species suggest that pollination function is restored along with pollinator abundance and richness; however, function may be less robust in restored habitats. An examination of interaction networks between bees and plant species found at both restored and remnant riparian sites showed less redundancy of pollinators visiting some plants at restored habitats.     

Recovery of Native Plant Community Characteristics on a Chronosequence of Restored Prairies Seeded into Pastures in West‐Central Iowa

Restored grasslands comprise an ever‐increasing proportion of grasslands in North America and elsewhere. However, floristic studies of restored grasslands indicate that our ability to restore plant communities is limited. Our goal was to assess the effectiveness of restoration seeding for recovery of key plant community components on former exotic, cool‐season pastures using a chronosequence of six restoration sites and three nearby remnant tallgrass prairie sites in West‐Central Iowa. We assessed trends in Simpson's diversity and evenness, richness and abundance of selected native and exotic plant guilds, and mean coefficient of conservatism (mean C). Simpson's diversity and evenness and perennial invasive species abundance all declined with restoration site age. As a group, restoration sites had greater richness of native C₃ species with late phenology, but lower richness and abundance of species with early phenology relative to remnant sites. Total native richness, total native abundance (cover), mean C, and abundance of late phenology C₃ plants were similar between restoration and remnant sites. Observed declines in diversity and evenness with restoration age reflect increases in C₄ grass abundance rather than absolute decreases in the abundance of perennial C₃ species. In contrast to other studies, restoration seeding appears to have led to successful establishment of tallgrass prairie species that were likely to be included in seeding mixtures. While several floristic measures indicate convergence of restoration and remnant sites, biodiversity may be further enhanced by including early phenology species in seeding mixes in proportion to their abundance on remnant prairies.     

High Plant Richness in Prairie Reconstructions Support Diverse Leafhopper Communities

A primary reason for restoring plant communities is to increase biodiversity to previous levels. It is expected that restoring land with greater plant diversity will increase biodiversity at higher trophic levels, but high diversity seed mixes are expensive. In this study, we used one insect family, leafhoppers (Hemiptera: Cicadellidae) to assess the difference in leafhopper communities that result from establishing high compared with low plant richness restorations. We tested the hypotheses that: (1) the added effort of a high richness restoration leads to measurable increases in both diversity and richness of leafhoppers; and (2) that leafhopper community composition is more similar to remnant prairies in high richness than in low plant richness restorations. We found that higher plant richness led to 3‐ to 7‐fold increases in leafhopper and prairie‐dependent leafhopper diversity and richness in restorations. Leafhopper communities in high richness restorations were not more similar to remnant prairies, rather they were distinct among high and low richness restorations and prairie interior. Leafhopper richness and diversity correlated with plant richness, and leafhopper community composition differed among plant community assemblages, but not with the occurrence of single plant species. For our sites, species‐rich restorations provided better quality habitat for leafhoppers that was comparable to remnant prairie. Our results suggest that restorations with high plant species richness better support animal food webs.     

Fruit‐feeding Butterfly Communities are Influenced by Restoration Age in Tropical Forests

Currently, a large‐scale restoration project aims to restore around 15 million hectares of Atlantic Forest in Brazil. This will increase forest cover and connectivity among remnant sites as well as restore environmental services. Currently, studies on recovery of fauna in restored areas of the Atlantic Forest are practically nonexistent. To address this knowledge vacuum, our study compares diversity patterns of fruit‐feeding butterflies in three forest areas with different restoration ages (11, 22, and 54 years), and uses a native forest area as reference. Results showed butterfly communities in maturing restored areas becoming more similar to the ones found in the native forest, with an increase in the proportional abundance of forest species, and a decrease of edge and grassland species. Moreover, we found a higher diversity among sites at the intermediate restoration age, with a community composed of both grassland and forest species. Butterfly species composition differed significantly among sites, showing interesting patterns of potential species replacement over time. Our results indicate that, although restored sites were located in a fragmented landscape, they provide suitable habitats for recolonization by fruit‐feeding butterfly assemblages. Hence, restored areas can be considered important habitat for forest animal species, increasing local biodiversity and, possibly, restoring some of the ecosystem services provided by them.     

Community structure of pollination webs of Mauritian heathland habitats

Pollination webs have recently deepened our understanding of complex ecosystem functions and the susceptibility of biotic networks to anthropogenic disturbances. Extensive mutualistic networks from tropical species-rich communities, however, are extremely scarce. We present fully quantitative pollination webs of two plant–pollinator communities of natural heathland sites, one of which was in the process of being restored, on the oceanic island of Mauritius. The web interaction data cover a full flowering season from September 2003 to March 2004 and include all flowering plant and their pollinator species. Pollination webs at both sites were dominated by a few super-abundant, disproportionately well-connected species, and many rare and specialised species. The webs differed greatly in size, reflecting higher plant and pollinator species richness and abundance at the restored site. About one fifth of plant species at the smaller community received <3 visits. The main pollinators were insects from diverse taxonomic groups, while the few vertebrate pollinator species were abundant and highly linked. The difference in plant community composition between sites appeared to strongly affect the associated pollinator community and interactions with native plant species. Low visitation rate to introduced plant species suggested little indirect competition for pollinators with native plant species. Overall, our results indicated that the community structure was highly complex in comparison to temperate heathland communities. We discuss the observed differences in plant linkage and pollinator diversity and abundance between the sites with respect to habitat restoration management and its influence on pollination web structure and complexity. For habitat restoration to be successful in the long term, practitioners should aim to maintain structural diversity to support a species-rich and abundant pollinator assemblage which ensures native plant reproduction.     

DNA metabarcoding uncovers dispersal-constrained arthropods in a highly fragmented restoration setting

Degraded areas are often restored through active revegetation; however, recolonization by animals is rarely engineered. Recolonization may be rapid for species with strong dispersal abilities. However, poor dispersers, such as many flightless arthropods, may struggle to recolonize newly restored sites. Actively reintroducing or “rewilding” arthropods may therefore be necessary to facilitate recolonization and restoration of arthropod communities and the ecological functions they perform. However, active interventions are rare. The purpose of this study was twofold. First, we asked whether potential source remnant arthropod communities were dispersal-constrained and struggling to recolonize restoration sites. Second, we tested whether reintroducing entire arthropod communities from remnant populations would help dispersal-constrained species establish during farmland ecological restoration in southern Australia. Rewilding was conducted in summer 2018 by transplanting leaf litter, soil, and entire communities contained within it from remnant source populations into geographically isolated restoration sites, which were paired with untreated controls (n = 6 remnant, rewilding transplant, and control sites). We collected leaf litter and extracted arthropod communities 19 months after the initial rewilding event, then sequenced mite, springtail, and insect communities using a metabarcoding approach. Within all groups, community similarity decreased with spatial distance between sites, suggesting significant dispersal barriers. However, only mite communities showed a strong response to rewilding, which was expressed as increased compositional similarity toward remnant sites and greater species richness relative to controls. Our results demonstrate that many arthropod species may struggle to recolonize geographically isolated restoration sites and that full community restoration requires active interventions via rewilding.     

Terrestrial Arthropods as Indicators of Ecological Restoration Success in Coastal Sage Scrub (California, U.S.A.)

Abstract Ecological restoration enjoys widespread use as a technique to mitigate for environmental damage. Success of a restoration project often is evaluated on the basis of plant cover only. Recovery of a native arthropod fauna is also important to achieve conservation goals. I sampled arthropod communities by pitfall trapping in undisturbed, disturbed, and restored coastal sage scrub habitats in southern California. I evaluated arthropod community composition, diversity, and abundance using summary statistics, cluster analysis, and detrended correspondence analysis (DCA) and investigated influence of vegetation on arthropod communities with multiple regression analysis. Arthropod diversity at undisturbed and disturbed sites was greater than at sites that were 5 and 15 years following restoration ( p < 0.05). Number of arthropod species was not significantly different among undisturbed, disturbed, and restored sites, and two restoration sites had significantly more individuals than other sites. Vegetation at disturbed and undisturbed sites differed significantly; older restorations did not differ significantly from undisturbed sites in diversity, percent cover, or structural complexity. In multiple regression models, arthropod species richness and diversity was negatively related to vegetation height but positively related to structural complexity at intermediate heights. Exotic arthropod species were negatively associated with overall arthropod diversity, with abundance of the earwig Forficula auricularia best predicting diversity at comparison (not restored) sites (r² = 0.29), and abundance of the spider Dysdera crocata and the ant Linepithema humile predicting diversity at all sites combined (r² = 0.48). Native scavengers were less abundant at restored sites than all other sites and, with a notable exception, native predators were less abundant as well. DCA of all species separated restored sites from all other sites on the first axis, which was highly correlated with arthropod diversity and exotic arthropod species abundance. Lower taxonomic levels showed similar but weaker patterns, with example families not discriminating between site histories. Vegetation characteristics did not differ significantly between the newly restored site and disturbed sites, or between mature restoration sites and undisturbed sites. In contrast, arthropod communities at all restored sites were, as a group, significantly different from both disturbed and undisturbed sites. As found in other studies of other restoration sites, arthropod communities are less diverse and have altered guild structure. If restoration is to be successful as compensatory mitigation, restoration success standards must be expanded to include arthropods.     

Initial soil amendments still affect plant community composition after nine years in succession on a heavy metal contaminated mountainside

Many efforts to restore disturbed landscapes seek to meet ecological goals over timescales from decades to centuries. It is thus crucial to know how different actions available to restoration practitioners may affect ecosystems in the long term, yet few such data exist. Here, we test the effects of seed and compost applications on plant community composition 9 years after their application, by taking advantage of a well‐controlled restoration experiment on a mountainside severely degraded by over 80 years of zinc smelting emissions. We asked whether plots have converged on similar plant communities regardless of initial seed and compost treatments, or if these initial treatments have given rise to lasting differences in whole plant communities or in the richness and abundance of native, exotic, and planted species. We found that compost types significantly affected plant communities 9 years later, but seed mix species composition did not. Observed differences in species richness and vegetative cover were negatively correlated, and both were related to the differences in plant communities associated with different compost types. These observed differences are due primarily to the number and abundance of species not in original seed mixes, of which notably many are native. Our results underscore the importance of soils in shaping the aboveground composition of ecosystems. Differences in soil characteristics can affect plant diversity and cover, which are both common restoration targets. Even in highly polluted and devegetated sites, compost and seed application can reinstate high vegetative cover and allow continued colonization of native species.     

Restoring species diversity: assessing capacity in the U.S. native plant industry

Large quantities of diverse and appropriately adapted native plant germplasm are required to facilitate restoration globally, yet shortages can prevent restorations from attaining desired species diversity and structure. An extensive native plant industry has developed in the United States to help meet these demands, yet very little is known about its capacity to support germplasm needs. To better understand current capacity and germplasm availability, we report results of the first comprehensive and quantitative assessment of the native plant industry in the United States, which includes at least 841 vendors nationwide and the species they make available for restoration. We synthesized lists of commercially available species from native plant vendors across the United States and identified gaps in species availability to inform germplasm research, development, and production. Of the approximately 25,000 vascular plant taxa native to the United States, 26% are sold commercially, with growth form, conservation status, distribution, and taxonomy significantly predicting availability. In contrast, only 0.07% of approximately 3,000 native nonvascular taxa are sold commercially. We also investigated how demand for germplasm to support high‐quality restoration efforts is met by vendors in the Midwestern tallgrass prairie region, which has been targeted extensively by restoration efforts for decades. In this well‐developed native plant market, 74% of more than 1,000 target species are commercially available, often from vendors that advertise genetically diverse, locally sourced germplasm. We make recommendations to build on the successes of regional markets like the tallgrass prairie region, and to fill identified gaps, including investing in research to support production, ensuring more consistent and clear demand, and fostering regional collaboration.     

Measuring Success: Evaluating the Restoration of a Grassy Eucalypt Woodland on the Cumberland Plain, Sydney, Australia

Abstract We compared the floristic composition and structure of restoration areas of eucalypt woodland with untreated pasture (control) and remnant vegetation (reference) in western Sydney. The restored areas comprised over 1,000 ha of abandoned pasture, which had been treated to reduce weeds and planted with seedlings of 26 native plant species raised from seed obtained locally from remnant vegetation. Plantings were carried out 0–9 years ago. Floristic composition was measured in quadrats using frequency scores and cover abundance. As far as possible treatments and restoration ages were replicated across sites. Ordination and analyses of similarity failed to distinguish the composition of restored vegetation from that of untreated pasture, which were both significantly different from that of remnant vegetation. There was a weak compositional trend with age of restored vegetation, but this was not in the direction of increasing resemblance to remnant vegetation. There was some evidence for convergence in structural features of restored with remnant vegetation, but this was at least partly attributed to plant growth. Subject to constraints imposed by the sampling design, environmental factors, and spatial variation were discounted as explanations for the results. The results therefore suggest either failure of restoration treatments or a restoration trajectory that is too slow to detect within 10 years of establishment. Our conclusions agree with those of similar studies in other ecosystems and support: (1) the need to monitor restoration projects against ecological criteria with rigorous sampling designs and analytical methods, (2) further development of restoration methods, and (3) regulatory approaches that seek to prevent damage to ecosystems rather than those predicated on replacing losses with reconstructed ecosystems.     

Beyond the green: assessing quarry restoration success through plant and beetle communities

In assessing the effectiveness of ecological restoration actions, outcomes evaluation using a multi‐taxa approach can greatly contribute to a clearer understanding of their success/failure. Since comprehensive biodiversity assessments are rarely possible, choosing taxa groups that are indicative of the ecosystem's structural and functional recovery is of major importance. Our goal was to evaluate the success of revegetation actions performed in a Mediterranean limestone quarry, using plants and epigean beetles as indicators. We compared their abundance, diversity, and community composition between revegetated sites aged 5, 13, and 19 years and a natural reference. Total plant cover significantly increased with restoration age and quickly reached reference values. However, native woody species cover dropped in the oldest site, while non‐native species became dominant. The abundance of beetles was always lower in restoration sites when compared to the reference, increasing with age, although not significantly. The richness of both plant species and beetle families was lower in restoration sites and did not show any trend towards the reference values. Finally, using nonmetric multidimensional scaling, the composition of plant and beetle communities from restoration sites showed a clear separation from the reference. Restoration efforts have successfully modified post‐quarry sites, but considerable differences remain, probably largely related to the use of the non‐native species Pinus halepensis in restoration plans. P. halepensis high cover in restoration sites greatly affects the structure of the ecosystem, and most likely its functioning too, as well as related ecosystem services, causing divergence from the reference values and compromising restoration success.     

Wild bee community change over a 26‐year chronosequence of restored tallgrass prairie

Restoration efforts often focus on plants, but additionally require the establishment and long‐term persistence of diverse groups of nontarget organisms, such as bees, for important ecosystem functions and meeting restoration goals. We investigated long‐term patterns in the response of bees to habitat restoration by sampling bee communities along a 26‐year chronosequence of restored tallgrass prairie in north‐central Illinois, U.S.A. Specifically, we examined how bee communities changed over time since restoration in terms of (1) abundance and richness, (2) community composition, and (3) the two components of beta diversity, one‐to‐one species replacement, and changes in species richness. Bee abundance and raw richness increased with restoration age from the low level of the pre‐restoration (agricultural) sites to the target level of the remnant prairie within the first 2–3 years after restoration, and these high levels were maintained throughout the entire restoration chronosequence. Bee community composition of the youngest restored sites differed from that of prairie remnants, but 5–7 years post‐restoration the community composition of restored prairie converged with that of remnants. Landscape context, particularly nearby wooded land, was found to affect abundance, rarefied richness, and community composition. Partitioning overall beta diversity between sites into species replacement and richness effects revealed that the main driver of community change over time was the gradual accumulation of species, rather than one‐to‐one species replacement. At the spatial and temporal scales we studied, we conclude that prairie restoration efforts targeting plants also successfully restore bee communities.     

Native soil microbial amendments generate trade‐offs in plant productivity,diversity, and soil stability in coastal dune restorations

Given the important role that soil microbes play in structuring plant communities and mediating ecosystem functions, there is growing interest in harnessing microbial communities to restore degraded ecosystems. Dune restorations, in particular, may benefit from native soil amendments because microbial diversity and abundance are very low in unvegetated areas. In an outdoor mesocosm experiment simulating Texas Gulf Coast dune restorations, we tested how native soil microbial amendments and restored diversity of foundational grasses influenced three key restoration responses: plant performance, plant diversity (including the colonization of native forbs), and soil stability. We found that native microbial amendments increased plant diversity and have the potential to increase soil stability, but this came at the cost of decreased plant biomass. Our results suggest that soil enemies in the native microbial amendments increased plant diversity by decreasing the performance of the dominant grass species and that arbuscular mycorrhizal fungi in the native microbial amendments increased the density of fungal hyphae in the soil, which can increase soil stability. Depending on the goals of the restoration, native soil microbial amendments may be a simple and inexpensive method to provide restoration benefits.     

Revegetation rewilds the soil bacterial microbiome of an old field

Ecological restoration is a globally important and well‐financed management intervention used to combat biodiversity declines and land degradation. Most restoration aims to increase biodiversity towards a reference state, but there are concerns that intended outcomes are not reached due to unsuccessful interventions and land‐use legacy issues. Monitoring biodiversity recovery is essential to measure success; however, most projects remain insufficiently monitored. Current field‐based methods are hard to standardize and are limited in their ability to assess important components of ecosystems, such as bacteria. High‐throughput amplicon sequencing of environmental DNA (metabarcoding of eDNA) has been proposed as a cost‐effective, scalable and uniform ecological monitoring solution, but its application in restoration remains largely untested. Here we show that metabarcoding of soil eDNA is effective at demonstrating the return of the native bacterial community in an old field following native plant revegetation. Bacterial composition shifted significantly after 8 years of revegetation, where younger sites were more similar to cleared sites and older sites were more similar to remnant stands. Revegetation of the native plant community strongly impacted on the belowground bacterial community, despite the revegetated sites having a long and dramatically altered land‐use history (i.e. >100 years grazing). We demonstrate that metabarcoding of eDNA provides an effective way of monitoring changes in bacterial communities that would otherwise go unchecked with conventional monitoring of restoration projects. With further development, awareness of microbial diversity in restoration has significant scope for improving the efficacy of restoration interventions more broadly.     

Passive restoration of vegetation and biological soil crusts following 80 years of exclusion from grazing across the Great Basin

Restoration targets for biological soil crusts are largely unknown. We surveyed seven 80‐year‐old grazing exclosures across northern Nevada for biocrusts to quantify reference conditions at relatively undisturbed sites. Exclosures were associated with the following plant communities: Wyoming big sagebrush, black sagebrush, and areas co‐dominated by winterfat and Wyoming big sagebrush. Cover of biocrusts and shrubs were generally higher than other plant groups at these sites, regardless of being inside or outside of the exclosures, suggesting these groups make up most of the native flora across the region. Important in forming soil structure, cyanobacteria of the order Oscillatoriales were less abundant and diverse in black sagebrush communities. Grazing had a negative effect on the abundance of Oscillatoriales but not the number of algal taxa, including cyanobacteria. Abundance of light algal crusts were not influenced by plant community or grazing. Dark algal crusts were generally less abundant on grazed sites. Influences of plant community and grazing were most apparent when accounting for reproductive rates of lichens and mosses based on establishment mechanisms. Abundance of shrubs, perennial grasses, Oscillatoriales, fast reproducing biocrusts and the number of algal and cyanobacterial taxa, varied by plant community, suggesting that restoration should be plant community specific. We demonstrate the affinity of rapidly reproducing biocrusts for winterfat‐Wyoming big sagebrush co‐dominated plant communities, regardless of grazing pressure. Across sites, the effects of grazing were most evident on the abundance of Oscillatoriales and slowly reproducing biocrusts following 80 years of cessation from grazing.     

Adaptation of plant‐mycorrhizal interactions to moisture availability in prairie restoration

The strength and direction of plant response to inoculation with arbuscular mycorrhizal fungi (AM fungi) is dependent on both abiotic and biotic contexts, often generating patterns of AM fungal mediation of plant adaptation. However, knowledge of plant‐community level effects of these interactions in grassland restoration is limited. We conducted a field inoculation experiment by inoculating five plant species native to a drier prairie and five plant species native to a moister prairie with mycorrhizal fungal communities from each prairie type. Species were paired by genus or family to account for phylogenetic effects. The inoculated plants were transplanted to study plots seeded with a restoration seed mix. Plots were manipulated to create either moister or drier conditions similar to environments of the plant species and mycorrhizal communities. In both transplanted and seeded plant species, we found that only drier prairie‐range species benefited from moisture‐regime matched AM fungal inoculum. Other seeded prairie plant species demonstrated a negative response to inoculation, likely due to the earlier successional stage of these species. Additionally, nonseeded plants benefited from inoculation in different ways: native nonseeded plants had highest cover with drier prairie inoculum in drier conditions, while nonnative plants had highest cover with moister prairie‐origin inoculum. These results suggest that use of local AM fungi may be particularly important in restorations at drier sites, even at relatively small differences in moisture availability. Further, specific knowledge of relative responsiveness of seeded plant species and nonseeded plant species to AM fungal inoculation will be useful in planning restorations.     

Species and functional trait re-assembly of ground beetle communities in restored grasslands

Ecosystem restoration provides unique opportunities to study community dynamics under succession and can reveal how consumer communities re-assemble and respond to successional changes. Studying community dynamics from both taxonomic and functional trait perspectives also may provide more robust assessments of restoration progress or success and allow cross-system comparisons. We studied ground beetle (Coleoptera: Carabidae) communities for three years in a restored grassland chronosequence with sites from 0 to 28 years old. We measured traditional community metrics (abundance, richness, Shannon diversity) and functional trait metrics based on species’ body length, wing morphology, activity time, phenology, and diet. Communities had high species richness and abundance in early successional stages, but these declined in later stages to low levels comparable to an adjacent grassland remnant. Species composition also shifted with time, converging with the remnant. Although functional richness, like species richness, declined as succession progressed, functional divergence quickly increased and was maintained over time, suggesting niche differentiation in established communities. Young sites were typified by small, macropterous, phytophagous species, while older sites contained larger species more likely to be flightless and carnivorous. Prescribed burns also affected traits, decreasing prevalence of larger species. This study demonstrates that functionally diverse consumer communities can self-assemble under restoration practices. In a relatively short amount of time both morphological and trophic level diversity are established. However, prescribed fire intended to control non-desirable plants may also shape beetle community functional composition, and restoration managers should consider if plant community benefits of fire outweigh potential declines in consumer function.     

Native-species seed additions do not shift restored prairie plant communities from exotic to native states

Exotic- and native-dominated communities can exist as alternate states in landscapes, but whether exotic-dominated states are persistent in the face of propagule pressure from native species is not well known. Here, we asked whether adding native seeds to low diversity, exotic-dominated patches would shift them to a more diverse, native state by using a long-term experiment with tallgrass prairie species in Iowa, USA. Previous work established that community assembly history led to alternate exotic or native states of perennial species. We added native seeds to plots in the spring after removing aboveground biomass with fire. We found that an experimental seed addition did not cause a shift from exotic to native states. Plots seeded eight years earlier in spring and without a priority effect continued to have the highest abundance and diversity of native species and lowest proportion of exotics. Our results suggest that exotic-dominated states in restorations can persist in the face of native species propagule pressure. Thus, assembly history can play a strong role in generating and maintaining alternate states over long time frames that are relevant to restoration. New restoration projects in exotic-dominated landscapes should maximize effort toward establishing native species during initial stages of restoration.     

Strengthening Invasion Filters to Reassemble Native Plant Communities: Soil Resources and Phenological Overlap

Preventing invasion by exotic species is one of the key goals of restoration, and community assembly theory provides testable predictions about native community attributes that will best resist invasion. For instance, resource availability and biotic interactions may represent “filters” that limit the success of potential invaders. Communities are predicted to resist invasion when they contain native species that are functionally similar to potential invaders; where phenology may be a key functional trait. Nutrient reduction is another common strategy for reducing invasion following native species restoration, because soil nitrogen (N) enrichment often facilitates invasion. Here, we focus on restoring the herbaceous community associated with coastal sage scrub vegetation in Southern California; these communities are often highly invaded, especially by exotic annual grasses that are notoriously challenging for restoration. We created experimental plant communities composed of the same 20 native species, but manipulated functional group abundance (according to growth form, phenology, and N‐fixation capacity) and soil N availability. We fertilized to increase N, and added carbon to reduce N via microbial N immobilization. We found that N reduction decreased exotic cover, and the most successful seed mix for reducing exotic abundance varied depending on the invader functional type. For instance, exotic annual grasses were least abundant when the native community was dominated by early active forbs, which matched the phenology of the exotic annual grasses. Our findings show that nutrient availability and the timing of biotic interactions are key filters that can be manipulated in restoration to prevent invasion and maximize native species recovery.