Restoration and Science: A Practitioner/Scientist's View from Rare Habitat Restoration at a Southern California Preserve

Researchers reexamining the relationship between restoration science and practice report a continuing scientist‐practitioner gap. As a land manager with scientific training, I offer my perspective of the chasm and describe a restoration practice infused with as much science as the realities of limited budget and time allow. The coastal sage scrub (CSS) restoration project at Starr Ranch, a 1,585 ha Audubon preserve in southern California, combines non‐chemical invasive species control, restoration, and applied research. Our practices evolve from modified scientific approaches and the scientific literature. Results from experiments with non‐optimum replication (on effects of seed rates, soil tamping, and timing of planting) nonetheless had value for management decisions. A critical practice came from academic research that encouraged cost‐effective passive restoration. Our passive restoration monitoring data showed 28–100% total native cover after 3–5 years. Another published study found that restoration success in semiarid regions is dependent on rainfall, a finding vital for understanding active restoration monitoring results that showed a range of 0–88% total native cover at the end of the first season. Work progresses through a combination of applied research, a watchful eye on the scientific literature, and “ecological intuition” informed by the scientific literature and our own findings. I suggest that it is less critical for academic scientists to address the basic questions on technique that are helpful to land managers but rather advocate practitioner training in methods to test alternative strategies and long‐term monitoring.     

Management of Non‐Native Annual Plants to Support Recovery of an Endangered Perennial Forb,Ambrosia pumila

Invasive non‐native plants pose a ubiquitous threat to native plant communities and have been blamed for the decline of many endangered species. Endangered species legislation provides legal instruments for protection, but identifying a general method for protecting endangered species by managing non‐natives is confounded by multiple factors. We compared non‐native management methods aimed at increasing populations of an endangered forb, Ambrosia pumila, and associated native plants. We compared the effects of a grass‐specific herbicide (Fusilade II), hand‐pulling, and mowing in two degraded coastal sage scrub sites in southern California, U.S.A. At both sites, hand‐pulling had the greatest effect on non‐native cover, and correspondingly resulted in the greatest increase in A. pumila stems. Fusilade II application also led to an increase in A. pumila, but was not as effective in controlling non‐native plants as hand‐pulling and its effect varied with the dominant non‐native species. Mowing was not effective at promoting A. pumila, and its effect on non‐native cover seemed to be related to rainfall patterns. Although some methods increased A. pumila, none of our treatments simultaneously increased cover of other native plants. Hand‐pulling, the most effective treatment, is labor intensive and thus only feasible at small spatial scales. At larger scales, managers should take an experimental approach to identifying the most appropriate method because this can vary depending on the specific management objective (endangered species or whole native community), the dominant non‐natives, yearly variation in weather, and the timing of treatment application.     

Native and naturalized plant diversity are positively correlated in scrub communities of California and Chile

Abstract. An emerging body of literature suggests that the richness of native and naturalized plant species are often positively correlated. It is unclear, however, whether this relationship is robust across spatial scales, and how a disturbance regime may affect it. Here, I examine the relationships of both richness and abundance between native and naturalized species of plants in two mediterranean scrub communities: coastal sage scrub (CSS) in California and xeric-sloped matorral (XSM) in Chile. In each vegetation type I surveyed multiple sites, where I identified vascular plant species and estimated their relative cover. Herbaceous species richness was higher in XSM, while cover of woody species was higher in CSS, where woody species have a strong impact upon herbaceous species. As there were few naturalized species with a woody growth form, the analyses performed here relate primarily to herbaceous species. Relationships between the herbaceous cover of native and naturalized species were not significant in CSS, but were nearly significant in XSM. The herbaceous species richness of native and naturalized plants were not significantly correlated on sites that had burned less than one year prior to sampling in CSS, and too few sites were available to examine this relationship in XSM. In post 1-year burn sites, however, herbaceous richness of native and naturalized species were positively correlated in both CSS and XSM. This relationship occurred at all spatial scales, from 400 m² to 1 m² plots. The consistency of this relationship in this study, together with its reported occurrence in the literature, suggests that this relationship may be general. Finally, the residuals from the correlations between native and naturalized species richness and cover, when plotted against site age (i.e. time since the last fire), show that richness and cover of naturalized species are strongly favoured on recently burned sites in XSM; this suggests that herbaceous species native to Chile are relatively poorly adapted to fire.     

High N,dry: Experimental nitrogen deposition exacerbates native shrub loss and nonnative plant invasion during extreme drought

Hotter, longer, and more frequent global change‐type drought events may profoundly impact terrestrial ecosystems by triggering widespread vegetation mortality. However, severe drought is only one component of global change, and ecological effects of drought may be compounded by other drivers, such as anthropogenic nitrogen (N) deposition and nonnative plant invasion. Elevated N deposition, for example, may reduce drought tolerance through increased plant productivity, thereby contributing to drought‐induced mortality. High N availability also often favors invasive, nonnative plant species, and the loss of woody vegetation due to drought may create a window of opportunity for these invaders. We investigated the effects of multiple levels of simulated N deposition on a Mediterranean‐type shrubland plant community in southern California from 2011 to 2016, a period coinciding with an extreme, multiyear drought in the region. We hypothesized that N addition would increase native shrub productivity, but that this would increase susceptibility to drought and result in increased shrub loss over time. We also predicted that N addition would favor nonnatives, especially annual grasses, leading to higher biomass and cover of these species. Consistent with these hypotheses, we found that high N availability increased native shrub canopy loss and mortality, likely due to the higher productivity and leaf area and reduced water‐use efficiency we observed in shrubs subject to N addition. As native shrub cover declined, we also observed a concomitant increase in cover and biomass of nonnative annuals, particularly under high levels of experimental N deposition. Together, these results suggest that the impacts of extended drought on shrubland ecosystems may be more severe under elevated N deposition, potentially contributing to the widespread loss of native woody species and vegetation‐type conversion.     

Cost‐effective ecological restoration

Ecological restoration is a multibillion dollar industry critical for improving degraded habitat. However, most restoration is conducted without clearly defined success measures or analysis of costs. Outcomes are influenced by environmental conditions that vary across space and time, yet such variation is rarely considered in restoration planning. Here, we present a cost‐effectiveness analysis of terrestrial restoration methods to determine how practitioners may restore the highest native plant cover per dollar spent. We recorded costs of 120 distinct methods and described success in terms of native versus non‐native plant germination, growth, cover, and density. We assessed effectiveness using a basic, commonly used metric (% native plant cover) and developed an index of cost‐effectiveness (% native cover per dollar spent on restoration). We then evaluated success of multiple methods, given environmental variation across topography and multiple years, and found that the most successful method for restoring high native plant cover is often different from the method that results in the largest area restored per dollar expended, given fixed mitigation budgets. Based on our results, we developed decision‐making trees to guide practitioners through established phases of restoration—site preparation, seeding and planting, and maintenance. We also highlight where additional research could inform restoration practice, such as improved seasonal weather forecasts optimizing allocation of funds in time or valuation practices that include costs of specific outcomes in the collection of in lieu fees.     

Grass invasion causes rapid increases in ecosystem carbon and nitrogen storage in a semiarid shrubland

Accurately predicting terrestrial carbon (C) and nitrogen (N) storage requires understanding how plant invasions alter cycling and storage. A common, highly successful type of plant invasion occurs when the invasive species is of a distinctly different functional type than the native dominant plant, such as shrub encroachment throughout the western United States and annual grass invasions in Mediterranean shrublands, as studied here. Such invasions can dramatically transform landscapes and have large potential to alter C and N cycling by influencing storage in multiple pools. We used a manipulation of non‐native annual grass litter within a shrub‐dominated habitat in southern California (coastal sage scrub, CSS) to study how grass invasion alters ecosystem C and N storage. We added, removed, or left unchanged grass litter in areas of high and low invasion, then followed soil and vegetation changes. Grass litter greatly increased C and N storage in soil, aboveground native and non‐native biomass. Aboveground litter storage increased due to the greater inputs and slower decomposition of grass litter relative to shrub litter; shading by grass litter further reduced decomposition of both non‐native and native litter, which may be due to reduced photodegradation. Soil C and N pools in areas of high litter increased ～20% relative to low litter areas in the two years following manipulation and were generally sinks for C and N, while areas with low litter were sources. We synthesize our results into a C cycle of invaded and uninvaded areas of CSS and link changes in storage to increases in the soil fungi : bacteria ratio, increased plant inputs, and decreased litter loss. Overall, we show that grasses, especially through their litter, control important abiotic and biotic mechanisms governing C and N storage, with widespread implications for C sequestration and N storage in semiarid systems undergoing grass or shrub invasions.     

Restoration of a Native Shrubland Impacted by Exotic Grasses, Frequent Fire, and Nitrogen Deposition in Southern California

Natural ecosystems globally are often subject to multiple human disturbances that are difficult to restore. A restoration experiment was done in an urban fragment of native coastal sage scrub vegetation in Riverside, California that has been subject to frequent fire, high anthropogenic nitrogen deposition, and invasion by Mediterranean annual weeds. Hand cultivation and grass‐specific herbicide were both successful in controlling exotic annual grasses and promoting establishment of seeded coastal sage scrub vegetation. There was no native seedbank left at this site after some 30 years of conversion to annual grassland, and the only native plants that germinated were the seeded shrubs, with the exception of one native summer annual. The city green‐waste mulch used in this study (C:N of 39:1) caused short‐term N immobilization but did not result in decreased grass density or increased native shrub establishment. Seeding native shrubs was successful in a wet year in this Mediterranean‐type climate but was unsuccessful in a dry year. An accidental spring fire did not burn first‐year shrubs, although adjacent plots dominated by annual grass did burn. The shrubs continued to exclude exotic grasses into the second growing season, suggesting that successful shrub establishment may reduce the frequency of the fire return interval.     

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.     

Vegetation-type conversion of evergreen chaparral shrublands to savannahs dominated by exotic annual herbs: causes and consequences for ecosystem function

Woody, evergreen shrublands are the archetypal community in mediterranean-type ecosystems, and these communities are profoundly changed when they undergo vegetation-type conversion (VTC) to become annual, herb-dominated communities. Recently, VTC has occurred throughout southern California chaparral shrublands, likely with changes in important ecosystem functions. The mechanisms that lead to VTC and subsequent changes to ecosystem processes are important to understand as they have regional and global implications for ecosystem services, climate change, land management, and policy. The main drivers of VTC are altered fire regimes, aridity, and anthropogenic disturbance. Some changes to ecosystem function are certain to occur with VTC, but their magnitudes are unclear, whereas other changes are unpredictable. I present two hypotheses: (1) VTC leads to warming that creates a positive feedback promoting additional VTC, and (2) altered nitrogen dynamics create negative feedbacks and promote an alternative stable state in which communities are dominated by herbs. The patterns described for California are mostly relevant to the other mediterranean-type shrublands of the globe, which are biodiversity hotspots and threatened by VTC. This review examines the extent and causes of VTC, ecosystem effects, and future research priorities.     

围封与放牧管理对高寒草甸植物功能性状和功能多样性的影响

植物功能性状能够响应生存环境的变化并直接决定着生态系统功能。为了揭示围封与放牧管理对物种共存和驱动群落构建的影响机理,该研究以青藏高原东缘高寒草甸为对象,分析了围封与放牧处理对植物功能性状和功能多样性的影响。结果显示:(1)在群落水平,放牧显著降低了比叶面积和植物高度;在物种水平,放牧群落中多数杂类草比叶面积减小,而莎草类和禾草类的比叶面积在处理间无显著差异。(2)叶干物质含量与比叶面积在围封和放牧处理中均呈显著负相关关系,在放牧处理中,叶干物质含量与植物高度呈显著的二次函数关系,即随着叶干物质含量的增大,植物高度先减小后增大;在同等比叶面积的情况下,与围封相比,放牧降低了叶干物质含量;在相同叶干物质含量的情况下,与围封相比,放牧降低了植物高度。(3)放牧在总体上降低了种间性状的平均差异,植物性状表现出趋同响应,具体表现为放牧减小了叶干物质含量和植物高度的种间差异;与围封相比,放牧显著提高了功能均匀度,减小了功能分离度。研究表明,不同植物种对放牧的响应模式存在差异,放牧降低了种间对光资源的竞争,可能增加了对土壤养分的竞争,放牧驱动群落构建的过程中,土壤养分是非常重要的作用因子,说明放牧影响物种共存依赖于对多种资源的竞争。     

Functional leaf and size traits determine the photosynthetic response of 10 dryland species to warming

Aims Relatively few studies so far have assessed how ongoing global warming will affect the photosynthetic performance of dryland plant species. We evaluated the effects of warming on the photosynthetic rates of 10 species with contrasting functional attributes, and whether their functional traits modulated photosynthetic responses to warming.Methods A common garden experiment was conducted over 2 years with distinct environmental conditions (drier vs. wetter year). The experiment was designed as a randomized block design with two treatments: warming (control vs. ~2.9°C temperature increase) and species (Agropyron cristatum, Festuca ovina, Lygeum spartum, Medicago sativa, Plantago lanceolata, Psoralea bituminosa, Sanguisorba minor, Hedysarum coronarium, Dorycnium pentaphyllum and Phlomis herba-venti). We linked functional traits measurements with temporal variations in photosynthetic responses to warming.Important findings In the drier year, warming increased photosynthetic rates at the beginning of the growing season, suggesting a modification in the growing period (earlier spring). In the wetter year, functional traits modulated photosynthetic responses to warming. Larger species with shorter leaves (e.g. M. sativa) had higher photosynthetic rates under warming compared to smaller species with larger leaves (e.g. F. ovina). Our results highlight the importance of (i) studying photosynthetic responses along different years and (ii) considering functional traits when evaluating photosynthetic responses to climate change, particularly in stressful environments such as drylands.     

Lake Restoration in New Zealand

Consideration of the need and opportunities for restoration of freshwater lakes in New Zealand requires individualised approaches that account for lake type, invasive species and food web structure.     

Exploring the potential of using priority effects during ecological restoration to resist biological invasions in the neotropics

Manipulating plant order of arrival, a process that creates priority effects, may be an unexplored powerful tool to hinder the establishment of invasive non‐native plants in sites under restoration. Knowledge and experimental studies on priority effects in the neotropics are scarce. Here, we propose a research agenda that investigates whether manipulating plant order of arrival can create priority effects in the neotropics, and if this strategy can be used to avoid the spread of undesired species in restoration projects. We also describe our view on expanding existing knowledge on priority effects to the neotropics and identifying key questions for future research.     

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.     

A trait-based approach across the native and invaded range to understand plant invasiveness and community impact

Alien plant species invasiveness and impact on diversity (i.e. species richness and composition) can be driven by the altered competitive interactions experienced by the invader in its invaded range compared to its native range. Trait-based competition effects on invasiveness can be mediated through size-asymmetric competition, i.e. a trait suit of the invader that drives competitive dominance, and through ‘niche differences', i.e. trait differentiation and thus minimized competition between invader and the invaded community. In terms of invasion impact, size-asymmetric competition is expected to result in competitive exclusion of co-occurring subordinate species, whereas ‘niche differences' might result in competitive exclusion of the most functionally similar co-occurring species. Although observational work does not allow the full disentanglement of both trait-based effects, it does allow to verify the occurrence of expected theoretical trait patters. In this study, we explored the trait-based competition effects on invasiveness and diversity impact for Rosa rugosa in both its invaded range in Belgium and its native range in Japan, based on seven functional traits across 100 vegetation plots. Following the predictions for enhanced invasiveness, we found much lower functional overlap between R. rugosa and the co-occurring species in the invaded range compared to the native range. This likely also explains the absence of diversity impact in its native range. Despite the absence of changes in species richness in the invaded range, the invader did strongly impact species composition of invaded communities. This impact occurred through strong shade tolerance responses, suggesting size-asymmetric competition effects and cover loss of co-occurring dominant species, next to exclusion of co-occurring species most functionally similar to the invader; suggesting niche difference effects. In conclusion, this case-study illustrates how exploring functional trait patterns across a species native and invaded range can help in understanding how trait-based competition processes can affect invasiveness and community impact.     

Competition between Native Hawaiian Plants and the Invasive Grass Megathyrsus maximus: Implications of Functional Diversity for Ecological Restoration

Biodiversity loss is a global crisis, due primarily to habitat destruction and widespread nonnative invasions. Invasive grasses are particularly problematic in many tropical ecosystems, where they possess traits that promote their persistence and can drastically alter native plant communities. We explored the ecophysiological basis for restoring native Hawaiian dryland ecosystems currently dominated by the nonnative invasive grass Megathyrsus maximus (guinea grass) in a garden experiment. Three native species—Myoporum sandwicense (naio; canopy tree), Dodonaea viscosa (aalii; shrub), and Plumbago zeylanica (iliee; groundcover)—were grown with M. maximus at three levels of native functional diversity (one, two, or three species) while holding overall plant density constant. We tested which individual and functional combinations of native species were more productive and best suppressed M. maximus growth and reproduction. Megathyrsus maximus had 39–94% higher maximum photosynthetic rates (A_max) than native species and increasing native functional diversity did not affect M. maximus A_max. Aboveground, belowground, and total biomass of M. maximus varied with functional diversity, although intraspecific competition reduced growth as much as interspecific competition. Reproductive tiller production by M. maximus decreased significantly when planted with any of the native species and with increasing native functional diversity. These results indicate that high native functional diversity in an ecological restoration setting may aid in the control of a dominant invasive grass and the reintroduction of diverse native species. Recommendations for restoring degraded nonnative grasslands in Hawaii and throughout the tropics include selection of native species that are ecophysiologically competitive and have high functional diversity.     

Functional diversity (FD), species richness and community composition

Functional diversity is an important component of biodiversity, yet in comparison to taxonomic diversity, methods of quantifying functional diversity are less well developed. Here, we propose a means for quantifying functional diversity that may be particularly useful for determining how functional diversity is related to ecosystem functioning. This measure of functional diversity “FD” is defined as the total branch length of a functional dendrogram. Various characteristics of FD make it preferable to other measures of functional diversity, such as the number of functional groups in a community. Simulating species' trait values illustrates how the relative importance of richness and composition for FD depends on the effective dimensionality of the trait space in which species separate. Fewer dimensions increase the importance of community composition and functional redundancy. More dimensions increase the importance of species richness and decreases functional redundancy. Clumping of species in trait space increases the relative importance of community composition. Five natural communities show remarkably similar relationships between FD and species richness.