Factoring restoration practitioner perceptions into future design of mechanical direct seeders for native seeds

The methods used to distribute seeds influence the success of a restoration project. We surveyed 183 restoration practitioners from across the globe with the aim of identifying common limitations to the effective use of mechanical direct seeding in large‐scale restoration practice to highlight avenues for design improvement to mechanized seeding equipment. Results from this survey show that direct seeding methods are commonly used for ecological restoration and agree with other studies that suggest the method can achieve results much quicker and cheaper than the alternative of distributing nursery‐grown tube stock. However, this study indicates that current mechanical direct seeding methods lack adequate control of seed sowing depth and spatial distribution and highlight that the inability to sow seeds of varying morphology over complex topography are common limitations to direct seeding. To improve restoration success, engineering improvements to mechanical direct seeders used in large‐scale restoration should focus in particular on addressing issues of precision of delivery for diverse seed types and landscapes.     

Restoring grassy woodland diversity through direct seeding: Insights from six ‘best‐practice’ case studies in southern Australia

Ecological restoration of grassy woodland ecosystems is now a significant landscape‐scale conservation objective throughout southern Australia. Technological improvements in direct seeding are now sufficiently well‐advanced to examine whether cost‐effective restoration of grassy woodlands is feasible. Consideration of six ‘best practice case studies shows substantial evidence of success. Further refinement of direct seeding techniques, in combination with native seed production systems, however, will be required into the future to meet the scale of woodland conservation targets and restore ecological function.     

Effectiveness of seed sowing techniques for sloped restoration sites

Practitioners are challenged with choosing among many potentially effective methods for sowing seed in ecological restoration projects to achieve sufficient native plant establishment. We tested the effectiveness of seed sowing techniques on moderate and steep slopes in a Mediterranean climate by measuring native seedling density immediately following germination, as well as plant density, recruitment success, and soil movement through the second growing season. We calculated cost effectiveness of different methods as the native plant density per dollar spent sowing seed. While all sowing techniques resulted in significant native establishment compared with unseeded controls, hydro seeding on moderate slopes was the most cost effective (native seedlings established per dollar spent). Although all steep‐sloped seeding techniques resulted in high densities of native species, all methods also resulted in significant soil loss. Shrubs preferred hand seeding followed by jute netting on steep slopes, while forbs reached greatest densities with hydro seeding on moderate slopes. Seedlings of species with heavy seeds were present in greater densities than species with lighter seeds in imprint sowing treatments. The “best” seed sowing technique varied depending on slope and metric of success (native density, species richness, shrub density, or forb density). Different combinations of slope, technique, and success metric resulted in significantly different project costs, which implies opportunities for savings given careful decision‐making relative to mitigation needs on heterogeneous landscapes. Evaluations of techniques for restoring slopes are limited, yet critical for expanding the area capable of being restored and the application of limited conservation funding.     

Using strip seeding to test how restoration design affects randomness of community assembly

The reestablishment and enhancement of plant diversity is typically a priority for restoration practitioners. Since diversity and stability can be affected by the magnitude to which randomness drives community dynamics, modifying randomness (via habitat heterogeneity) could provide utility for vegetation managers. We investigated the value of using strip seeding to manipulate the magnitude to which randomness structures plant communities across a grassland in Davis, California. Five years after restoring portions of a degraded site (0, 33, 50, 66, and 100% of an area) to create patches of seeded and unseeded strips, we assessed the amount of Jaccard dissimilarity across quadrats within strips and estimated the magnitude to which randomness contributed to community assembly (termed the nugget). We found higher nuggets in the 66 and 33% seeding treatment levels compared to the 0, 50, and 100% seeding treatment levels. In the 33 and 66% level of the seeding treatment, we also found that unseeded strips, which are regularly exposed to random events of dispersal from seeded strips, had a higher nugget than seeded strips. This work suggests that strategic seeding techniques that enhance habitat heterogeneity can increase the role of randomness in community dynamics. Strip seeding strategies appear to provide utility as a tool to indirectly enhance diversity across a degraded site.     

Seed use in the field: delivering seeds for restoration success

Seed delivery to site is a critical step in seed‐based restoration programs. Months or years of seed collection, conditioning, storage, and cultivation can be wasted if seeding operations are not carefully planned, well executed, and draw upon best available knowledge and experience. Although diverse restoration scenarios present different challenges and require different approaches, there are common elements that apply to most ecosystems and regions. A seeding plan sets the timeline and details all operations from site treatments through seed delivery and subsequent monitoring. The plan draws on site evaluation data (e.g. topography, hydrology, climate, soil types, weed pressure, reference site characteristics), the ecology and biology of the seed mix components (e.g. germination requirements, seed morphology) and seed quality information (e.g. seed purity, viability, and dormancy). Plan elements include: (1) Site treatments and seedbed preparation to remove undesirable vegetation, including sources in the soil seed bank; change hydrology and soil properties (e.g. stability, water holding capacity, nutrient status); and create favorable conditions for seed germination and establishment. (2) Seeding requirements to prepare seeds for sowing and determine appropriate seeding dates and rates. (3) Seed delivery techniques and equipment for precision seed delivery, including placement of seeds in germination‐promotive microsites at the optimal season for germination and establishment. (4) A monitoring program and adaptive management to document initial emergence, seedling establishment, and plant community development and conduct additional sowing or adaptive management interventions, if warranted. (5) Communication of results to inform future seeding decisions and share knowledge for seed‐based ecological restoration.     

Direct seeding and outplantings in drylands of Argentinean Patagonia: estimated costs,and prospects for large‐scale restoration and rehabilitation

In large areas of the world that are deeply scarred by desertification and hampered by low capacity for natural regeneration, the scaling up of ecological restoration and rehabilitation can be achieved only if it is low in cost with high return on investment, and shows promise of providing long‐lasting social‐economic as well as ecological benefits. In the Monte Austral region of Patagonia Argentina, concerted efforts are underway to facilitate scaling up of ecological restoration and rehabilitation practices. Here, we evaluate financial costs and preliminary results of direct seeding as compared to outplanting of nursery‐grown seedlings of three native species (Atriplex lampa, Senecio subulatus var. subulatus, and Hyalis argentea var. latisquama) considered to be high‐priority dryland framework species. Comparative success is expressed in terms of plant survival and in monetary terms. The three candidate species showed low survival rates, ranging from 4.3 to 22.3%, after the first summer following direct seeding. In contrast, survival rates for planted seedlings of the same three taxa varied between 84 and 91%, after the first summer following reintroduction. However, cost of direct seeding varied between 1,693 and 1,772 US$ less per hectare, that is, 64% less than the cost of outplanting nursery seedlings. Therefore, in the search for ways to scale up ecological restoration and rehabilitation in drylands, direct seeding should receive more attention. We discuss the social and ecological perspectives and the way forward for direct seeding techniques in Patagonia. We also consider how costs could be reduced and effectiveness improved in large‐scale efforts.     

Diminishing Returns from Higher Density Restoration Seedings Suggest Trade‐offs in Pollinator Seed Mixes

Native forbs have become a more central component of restoration programs, especially because of their role in supporting crop pollinators. This study evaluates the success of different native forb mixes and seeding rates using shared goals of restoration practitioners and agroecologists, namely percent native species cover, floral resources, native diversity, and cost‐effectiveness. At 6 sites with hedgerows adjacent to agricultural lands in California's Central Valley, we planted 3 native forb seed mixes at 3 seeding rates and monitored germination, percent cover, and floral resources for 2 to 3 years. We also evaluated the cost of the mixes based on seeding rates and original seed prices. More than mix type, relative seeding rate strongly affected germination, cover, and floral resource success. The relative benefits of seeding with more species diminished at higher seeding rates, especially when cost was considered. Cover increased significantly over the years but diversity declined sharply after the first year. Increased cover of target species was mainly due to the effect of 1 dominant species Grindelia camporum, common gumplant. We summarize data from a similar forb restoration study showing that the species that dominated in our mix‐and‐rate experimental sites also attracted the greatest diversity and abundance of pollinators. These findings highlight trade‐offs and balance‐points within restoration and pollination services goals. We offer suggestions on how to weigh those trade‐offs, given particular priorities and how native forb plantings can support combined goals of pollination services and restoration.     

Seedling Establishment and Survival on Restored Campsites in Subalpine Forest

This study experimented with common restoration techniques (scarification, soil amendments, mulch, and seeding) on six closed wilderness campsites in subalpine forests in Oregon. Effectiveness in encouraging seedling establishment, growth, and survival was assessed every year for the first 7 years following treatment. Closure and restoration of the campsites increased the density of plants established from seed. Despite an original density of virtually zero, mean density of perennial plants was 55 plants/m² 7 years after closure. All the treatments, with the exception of the biodegradable mulch mat, increased plant density. Seven years after treatment, seeding had increased plant density 5‐fold, whereas scarification and soil amendments (organic matter, compost, and soil inoculum) had each increased density 3‐fold. The organic and compost amendments also had the positive benefit of increasing growth rates and shortening the time‐to‐reproductive maturity. Results suggest that restoration of the herbaceous cover on these campsites can occur rapidly using the techniques employed. All but one of the species we seeded established in substantial quantities and survived at densities exceeding their density in the naturally sparse herbaceous cover on these sites. Thirty‐six perennial species volunteered on these sites. The remaining challenge is reestablishment of the shrub species that comprise much of the ground cover in these forests. These species seldom establish from seed.     

Recovery after African Olive invasion: can a ‘bottom‐up’ approach to ecological restoration work?

African Olive (Olea europaea ssp. cuspidata) is a densely crowned evergreen small tree, native to eastern Africa that is highly invasive in areas where it has been introduced, including Hawaii and Australia. Invasion by African Olive threatens Cumberland Plain Woodland, a critically endangered grassy eucalypt woodland from western Sydney, Australia, through the formation of a dense mid‐canopy excluding the regeneration of native species. We established a 3‐year field experiment to determine the effectiveness of direct seeding and fire, as techniques for early stage restoration of a 2 ha historically cleared and degraded Cumberland Plain Woodland site after the removal of African Olive. Direct seeding was able to re‐establish a native perennial grass cover which was resistant to subsequent weed invasion and could be managed as an important first stage in woodland restoration with fire and selective herbicide. Fire was able to stimulate some germination of colonising native species from the soil seed bank after 15 years of African Olive invasion; however, germination and establishment of native shrubs from the applied seed mix was poor. We propose a ‘bottom‐up’ model of ecological restoration in such highly degraded sites that uses a combination of direct seeding and stimulation of the soil seed bank by fire, which could be applicable to other degraded grassy woodland sites and plant communities.     

Trait‐based approach confirms the importance of propagule limitation and assembly rules in old‐field restoration

Community assembly theory is suggested as a guiding principle for ecological restoration to help understand the mechanisms that structure biological communities and identify where restoration interventions are needed. We studied three hypotheses related to propagule limitation, stress‐dominance, and limiting similarity concepts in community assembly in a restoration field experiment with a trait‐based null model approach. The experiment aimed to assist the recovery of sand grassland on former arable land in the Kiskunság, Pannonian biogeographic region, Europe. Treatments included initial seeding of five grassland species, carbon amendment, low‐intensity mowing, and combinations in 1 m by 1 m plots in three old fields from 2003 to 2008. The distribution of 10 individual plant traits was compared to the null model and the effect of time and treatments were tested with linear mixed effect models. Initial seeding had the most visible impact on species and trait composition confirming propagule limitation in grassland recovery. Reducing nutrient availability through carbon amendment strengthened trait convergence for length of flowering as expected based on the stress‐dominance hypothesis. Mowing changed trait divergence to convergence for plant height with a strengthening impact with time, supporting our hypothesis of increasing dominance of limiting similarity with time. Our results support the idea that community assembly is simultaneously influenced by propagule limitation and multiple trait‐based processes that act through different traits. The limited impact of manipulating environmental filtering and limiting similarity compared to seeding, however, supports the view that only targeting the dispersal and environmental filters in parallel would improve restoration outcome.     

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.     

Should I plant or should I sow? Restoration outcomes compared across seven riparian revegetation projects

To revegetate native plant communities, it is often cheaper to direct seed than to plant nursery‐grown stock. However, the outcomes of direct seeding can be quite variable, and it is unclear whether direct seeding or planting is more likely to facilitate the restoration of diverse plant communities. To address this question, we compared the outcomes of each method across several recent riparian revegetation projects where both direct seeding and tube‐stock planting were used. We surveyed riparian revegetation projects at seven sites within the greater Melbourne area that had been revegetated between 1 and 4 years previously. Sites were all on land previously used for agriculture or degraded public land and ranged in environmental and climatic conditions. Woody plant density, establishment of target species, species richness, species diversity (evenness) and plant heights were assessed. Direct seeding tended to result in higher plant densities and similar species richness, but lower rates of species establishment and diversity compared with planting. A median of 67% of target species established via direct seeding compared with 100% for planting, with direct seeded areas often dominated by one or two species. In general, overall revegetation outcomes were often driven by climatic and site factors, rather than revegetation method. We suggest that to achieve good restoration outcomes from revegetation in riparian areas, a bet‐hedging or combined approach using both sowing and planting may be the best strategy.     

Expanding horizons for herbaceous ecosystem restoration: the Grassy Groundcover Restoration Project

Summary The Grassy Groundcover Restoration Project (GGRP) has sown thirteen 1 ha plots of species‐rich grassland or herbaceous understorey in previously weedy agricultural paddocks in a range of rural locations across southern and western Victoria, Australia. The sown plots are intended as both experimental trials and ‘core’ areas for the restoration of herbaceous communities native to these regions. Approximately 200 species were grown in Seed Production Areas (SPAs) and successfully sown in the field. Species were most successfully established on areas that were scalped prior to seeding, and least successful on plots that were pre‐treated with 1, 2 or 3 years of traditional herbicide weed control. Weed presence was lowest in scalped plots and highest in non‐scalped plots. Long‐term monitoring will be required to understand the development trajectories and degree of persistence of the sown communities, but in the shorter term (3–6 years of post‐seeding) an average of 80% of sown species have established and remain as adult populations. Surveys indicate that in scalped plots (n = 130) vegetation composition, structure and quality has been maintained. Conversely, composition, structure and quality have declined markedly in non‐scalped plots (n = 130). Formal surveys and field observations have also revealed that all sites provide a range of habitats which have been colonized by fauna from a variety of trophic levels. The implications of building on these trials to realize complex grassy ecosystem restoration at larger scales are discussed including the securing of sufficient quantities of high‐quality seed, the use of mechanized broad‐scale direct‐seeding techniques and the effectiveness of using complex mixtures of species early in the restoration cycle.     

Optimizing seeding density of fast‐growing native trees for restoring the Brazilian Atlantic Forest

Direct seeding is a promising method for reducing restoration costs, but methodological adjustments are still needed to reduce the uncertainties to achieve a desired seedling density in the field. Here, we investigated the technical approaches and outcomes of direct seeding of fast‐growing native trees for cost‐effective restoration of the Brazilian Atlantic Forest. Sixteen tree species were manually sown at three seeding densities in planting lines prepared with a subsoiler, in two experimental areas, which were weeded with hoes and had leaf‐cutter ants controlled with insecticide baits. Seedling density was monitored for 30, 90, and 180 days after sowing. No substantial change in tree density was observed 30 days after sowing, thus allowing fast corrective actions to adjust tree density. Only a minor proportion of the sown viable seeds resulted in established seedlings at 180 days (4–12% for the community; approximately 25% for the species with the best performance). However, tree density was high (6,000 on average; approximately 1,400–13,000 trees/ha) and allowed an effective canopy development. Overall, seedling density was linearly and positively associated with seeding density, was highly influenced by the species used, and was higher in the soil with higher sum of bases. Buying seeds would be, for most species, less costly than buying nursery‐grown seedlings for achieving the expected tree densities in the field. These results evidence the potential of direct seeding for reducing restoration costs, as well as the need to select species with better performance and adjust seeding densities to optimize the use of this method.     

Adapting management to a changing world: Warm temperatures,dry soil,and interannual variability limit restoration success of a dominant woody shrub in temperate drylands

Restoration and rehabilitation of native vegetation in dryland ecosystems, which encompass over 40% of terrestrial ecosystems, is a common challenge that continues to grow as wildfire and biological invasions transform dryland plant communities. The difficulty in part stems from low and variable precipitation, combined with limited understanding about how weather conditions influence restoration outcomes, and increasing recognition that one‐time seeding approaches can fail if they do not occur during appropriate plant establishment conditions. The sagebrush biome, which once covered over 620,000 km² of western North America, is a prime example of a pressing dryland restoration challenge for which restoration success has been variable. We analyzed field data on Artemisia tridentata (big sagebrush) restoration collected at 771 plots in 177 wildfire sites across its western range, and used process‐based ecohydrological modeling to identify factors leading to its establishment. Our results indicate big sagebrush occurrence is most strongly associated with relatively cool temperatures and wet soils in the first spring after seeding. In particular, the amount of winter snowpack, but not total precipitation, helped explain the availability of spring soil moisture and restoration success. We also find considerable interannual variability in the probability of sagebrush establishment. Adaptive management strategies that target seeding during cool, wet years or mitigate effects of variability through repeated seeding may improve the likelihood of successful restoration in dryland ecosystems. Given consistent projections of increasing temperatures, declining snowpack, and increasing weather variability throughout midlatitude drylands, weather‐centric adaptive management approaches to restoration will be increasingly important for dryland restoration success.     

Are two strategies better than one? Manipulation of seed density and soil community in an experimental prairie restoration

Restoration practitioners have a variety of practices to choose from when designing a restoration, and different strategies may address different goals. Knowledge of how to best use multiple strategies could improve restoration outcomes. Here, we examine two commonly suggested strategies in a single tallgrass prairie restoration experiment: increased forb sowing density and prairie soil inoculation. We designed a study with two different forb seeding densities. Within these densities, we transplanted seedlings into 1‐m² plots that had been grown in either a whole prairie soil inoculum or sterilized prairie soil. After 4 years, we found positive effects of both high forb sowing density and inoculation treatments on the ratio of seeded to nonseeded plant cover in these plots, and negative effects of both treatments on nonseeded plant diversity. No effects of either treatment were seen on seeded plant diversity. Each strategy also affected the plant community in different ways: high forb sowing density increased seeded forb richness and decreased native nonseeded plant cover, while inoculation decreased non‐native cover, and tended to increase average successional stage of the community. These effects on restoration outcome were typically independent of each other, with the result that plots with both manipulations had the most positive effects on restoration outcomes. We thus advocate the combined use of these restoration strategies, and further studies which focus on both seeding and soil community manipulation in restoration.     

The early response of subtropical tussock grasslands to restoration treatments

Once widespread, Australia's bluegrass tussock grasslands (dominated by Dichanthium sp.) of the Queensland Central Highlands are now severely endangered. Despite being biodiversity rich and highly valued as low input, nutrient‐dense grazing systems, bluegrass tussock grasslands have suffered extensive clearing and degradation over the last 150 years. Natural recovery of these grasslands is possible but rates of recovery are slow. As such, there is an urgent need to assess practical management strategies to accelerate recovery of these grasslands, with a particular focus on early‐successional stages, when aggressive exotic species are most prevalent. To date, no studies have tested whether commonly used grassland restoration strategies can enhance early‐successional stages and accelerate regeneration in this system. Here, we examine the early short‐term impacts (first two seasons) of two common grassland restoration approaches, with two variations each: direct seeding (single species and low seed diversity) and vegetation clearing (prescribed burning and glyphosate application) across two common starting points: a formerly cropped old field and a historically overgrazed natural grassland. No treatment increased native diversity (Shannon's or richness) in plots but the composition of burned plots in the old field did become more similar to healthy reference sites after two seasons. Burning combined with direct seeding also increased the abundance of the dominant grass, Dichanthium sericeum, toward healthy reference levels within the first two seasons post seeding. This study provides a practical assessment of the short‐term impacts and capacity of common grassland restoration treatments to enhance the recovery of Australia's tussock grassland systems.     

Long‐term effects of prairie restoration on plant community structure and native population dynamics

The key to restoring degraded grassland habitats is identifying feasible and effective techniques to reduce the negative impacts of exotic species and promote self‐sustaining native populations. It is often difficult to extend monitoring of restoration efforts to evaluate long‐term success, but doing so is essential to understanding how initial outcomes change over time. To assess how initial treatment effects persist, we revisited degraded patches of Pacific Northwest prairie habitat 6 years after experimental restoration efforts ceased. We evaluated plant community composition to determine the lasting effects of supplemental native seeding and disturbance treatments (burning, mowing, and herbicide to reduce exotic species). We tracked the persistence of seeded species and measured spread of their populations to evaluate suitability of species for restoration and the ability of the habitat to support native plant populations. We found that plots that received supplemental seeding continued to exhibit higher richness of native species than those left unseeded, and that both seeding and disturbance treatments could positively influence native species abundance over the long term. The initially observed effects of disturbance treatments on reducing exotic grass abundance had diminished, highlighting the importance of long‐term monitoring and ongoing control of exotic species. Nevertheless, these treatments significantly influenced the population trajectories of 4 out of 8 seeded native species. There was evidence of spatial advance of most seeded species. Results from extended monitoring confirm that dispersal limitation of native species and difficulties maintaining the reduction of exotic grasses continue to be major barriers to success in restoration of invaded grasslands.     

Restoration,soil organisms,and soil processes: emerging approaches

Better understanding of the connection between aboveground plant communities and belowground soil organisms and processes has led to an explosion in recent research on the applications of this link to the field of ecological restoration. Research is only beginning to have the capacity to link soil organisms and specific ecosystem functions. Establishing general ecological principles of the role microbial communities have during ecological restoration is also still in its infancy. As such, the literature is at a critical point to generate a Special Feature that brings together novel approaches of linking soil and restoration to promote more regular inclusion and consideration of soil organisms and soil‐based processes in ecological restoration. In this special feature, we bring together nine research articles from different ecosystems that study the relationship between restoration activities, soil microbial communities, and soil properties. From these research articles, we describe two primary themes: (1) research on the impacts of ecosystem‐specific restoration activities on soil organisms and processes and (2) research testing methods of soil manipulation to improve restoration outcomes. We hope to inspire readers and restoration practitioners to consider soil microbes and soil processes in their research, restoration projects, and world views.     

Does Seeding After Wildfires in Rangelands Reduce Erosion or Invasive Species?

Mitigation of ecological damage caused by rangeland wildfires has historically been an issue restricted to the western United States. It has focused on conservation of ecosystem function through reducing soil erosion and spread of invasive plants. Effectiveness of mitigation treatments has been debated recently. We reviewed recent literature to conduct a meta‐analysis of seeding after wildfires to determine if seedings may (1) protect ecosystems against soil erosion and (2) reduce invasion or abundance of undesirable nonnative plant species. Effectiveness of postfire seedings was examined in 8 erosion and 19 invasive species cases. Seeding has little effect on erosion during the first year after fire and is highly dependent upon initial establishment and coverage of species in successive years. Among all seeding cases, 28% reduced, 67% were neutral, and 5% increased invasive species abundance. Older seedings were more likely to show reductions in invasives than younger seedings. Seedings with high plant establishment were more likely to reduce invasives than those with low establishment. Studies are needed that examine (1) frequency of adequate establishment of postfire seedings and causal factors of success or failure, (2) long‐term impacts of seeding along a range of initial establishment and concomitant plant coverage over time as it relates to erosion and abundance of invasive plant species, and (3) auxiliary treatments designed to increase likelihood of germination and establishment given the inevitable variability of environmental conditions. These studies would aid land managers in deciding when postfire treatments are required and their likely level of success.