Restoring the Victorian Western (Basalt) Plains grassland. 1. Laboratory trials of viability and germination, and the implications for direct seeding

Summary   The Victorian Western (Basalt) Plains grassland is one of Australia's most threatened plant communities. Practitioners using seed for its restoration need to know whether seed can be sown fresh or whether it requires an after-ripening period. This study assessed the viability and germination of freshly harvested wild seed from 64 grassland species indigenous to the Basalt Plains of western Victoria. The seed was collected as part of a broader experiment that examined the potential of direct-sown complex seed mixes for the restoration of grassland communities. The germination of fresh seed at 25°C varied widely between species. Comparisons with tetrazolium viability tests for each species indicated varying levels of dormancy within the species pool. Germination separated into three broad responses at day 28. One-third of the species failed to germinate, one-third germinated at 1% to 50% and the remaining species germinated between 51% and 100%. Therefore, if the aim of a sowing was the rapid and synchronous establishment of most of the sown species, the use of fresh seed in restoration could be problematic. After 3 months of dry storage, eight species were re-tested for germination. Each of the selected species had shown high viability but low initial germination. Only two species significantly increased their total germination at 25°C. The annual species, Triptilodiscus pygmaeus , increased its total germination from 6% as fresh seed to 99% after dry storage. Testing the viability and germination capacity of freshly harvested seed from a large and diverse sample of native grassland species demonstrated that many of the species were unlikely to germinate rapidly or synchronously when sown in complex seed mixes soon after harvest. This finding has implications for the scheduling and management of restoration projects that rely on the use of such seed.     

Effect of artificial shade and grazing removal on degraded grasslands: Implications of woody restoration for herbaceous vegetation

Extensive degraded short tussock grasslands of New Zealand's eastern South Island were dominated by woody vegetation prior to burning and livestock grazing associated with human settlement starting 800 years ago. There is increasing interest in restoring some of these grasslands back to a woody state. However, because of the long time frames involved in establishing a woody cover, it is difficult to predict the impacts that woody restoration will have on the extant herbaceous flora. Using a factorial trial with artificial shade and grazing exclusion, we assessed the potential impact of woody restoration on the structure and composition of the herbaceous flora over a six‐year period. The imposition of artificial shade resulted in significant increases in total species richness and the total cover of herbaceous vegetation, increases in cover of several individual forb and grass species and decreases in the cover of bare ground, moss and lichen in shade treatments. There were also changes in the overall community composition of shaded treatments reflecting these changes in vegetation cover and species richness. We found no statistically significant effects of grazing exclusion. We suggest that increased soil moisture resulting from shade addition plays an important role in increasing the herbaceous component of the flora. While woody restoration will have a range of effects on the herbaceous understorey, for example through competition and changes in soil conditions, our findings are important for planning future woody restoration in these degraded tussock grasslands. In particular, our results suggest that the best approach to ensure the persistence of herbaceous vegetation in woody restorations might be to ensure that restoration plantings result in a spatially heterogeneous vegetation arrangement.     

Reconstructing grassy understories in south‐eastern Australia: Interview with Paul Gibson‐Roy

A marriage of ecology with agronomy is proving successful in restoring diverse herbaceous layers ‐ to the extent that some reconstructed grasslands on ex‐agricultural land and rural roadsides have been found eligible for federal protection as threatened ecological communities. Can lessons from this improve our management and expansion of grassy ecosystems more broadly?     

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.     

Seeding the future – the issues of supply and demand in restoration in Australia

It has been almost 15 years since concerns about the limited capacity of remnant native vegetation to supply the volumes of seed required to meet increasing restoration demands were first raised. Since that time little progress has been made towards addressing this constraint with the ongoing decline of native vegetation communities, especially since 2000, further challenging seed supply. We provide examples of the size of this demand for seed, as well as major issues associated with seed sourcing. We also discuss how invoking the concept of market forces to drive seed supply and demand is inappropriate and highlight the need for an industry body to oversee seed collection and utilisation standards. We further propose key actions that are required to secure the seed supply chain within the next 20 years to meet existing and future restoration targets. We argue that concerted, coordinated action at Commonwealth, State and regional levels are required to underpin effective future restoration outcomes.     

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.     

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.     

Predicting the establishment success of introduced target species in grassland restoration by functional traits

Species‐rich semi‐natural grasslands are highly endangered habitats in Central Europe and numerous restoration efforts have been made to compensate for the losses in the last decades. However, some plant species could become more easily established than others. The establishment success of 37 species was analyzed over 6 years at two study sites of a restoration project in Germany where hay transfer and sowing of threshing material in combination with additional sowing were applied. The effects of the restoration method applied, time since the restoration took place, traits related to germination, dispersal, and reproduction, and combinations of these traits on the establishment were analyzed. While the specific restoration method of how seeds were transferred played a subordinate role, the establishment success depended in particular on traits such as flower season or the lifeform. Species flowering in autumn, such as Pastinaca sativa and Serratula tinctoria, became established better than species flowering in other seasons, probably because they could complete their life cycle, resulting in increasingly stronger seed pressure with time. Geophytes, like Allium angulosum and Galium boreale, became established very poorly, but showed an increase with study duration. For various traits, we found significant trait by method and trait by year interactions, indicating that different traits promoted establishment under different conditions. Using a multi‐model approach, we tested whether traits acted in combination. For the first years and the last year, we found that models with three traits explained establishment success better than models with a single trait or two traits. While traits had only an additive effect on the establishment success in the first years, trait interactions became important thereafter. The most important trait was the season of flowering, which occurred in all best models from the third year onwards. Overall, our approach revealed the potential of functional trait analysis to predict success in restoration projects.     

Establishment of tree seedlings in the understory of restoration plantations: natural regeneration and enrichment plantings

Little is known about the potential of restoration plantations to provide appropriate understory conditions to support the establishment of seeds arriving from neighboring native forests. In this article, we investigated how seedling establishment is affected in the understory of restoration sites of different ages and assessed some of the potential environmental factors controlling this ecological process. We first compared the density and richness of native tree seedlings among 10‐, 22‐, and 55‐year‐old restoration plantations within the Atlantic Forest region of southeastern Brazil. Then, we undertook a seed addition experiment in each study site, during the wet season, and compared seedling emergence, survival, and biomass on local versus old‐growth forest soil (transferred from a reference ecosystem), in order to test whether local substrate could hamper seedling establishment. As expected, the oldest restoration site had higher density and richness of spontaneously regenerating seedlings. However, seedling establishment was less successful both in the oldest restoration planting and using substrate transferred from a reference ecosystem, where emergence and survival were lower, but surviving seedlings grew better. We attribute these results to lower light availability for seedlings in the understory of the oldest site and speculate that higher incidence of pathogens on old‐growth forest soil may have increased seedling mortality. We conclude that the understory of young restoration plantations provides suitable microsite conditions at the early establishment phases for the spontaneous regeneration or enrichment planting of native trees.     

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.     

Restoration Ecology's silver jubilee: innovation,debate, and creating a future for restoration ecology

At a time when the science and practice of restoration ecology is adapting to ongoing environmental and social change, innovations in both methods and concepts are essential. Encouraging innovation means allowing open debate about alternative approaches that may add to the toolbox available for restoration. Such approaches are usually being examined as additions to, rather than substitutes for, traditional restoration practices. Recent debate has focused on the scope and intent of restoration as defined in documents such as the Society for Ecological Restoration Standards. There is a mismatch between the default aim in the standards of full restoration to a native reference system and the goals of international restoration efforts that have a broader and more functional focus. The next generation of restoration scientists and practitioners will need to navigate these issues to ensure that restoration remains effective and relevant. This will require, amongst other things, ongoing learning, sharing information and insights, humility, objectivity, continuous examination of assumptions, and questioning current practices and perspectives.     

A world of possibilities: six restoration strategies to support the United Nation's Decade on Ecosystem Restoration

Ecological restoration is practiced worldwide as a direct response to the degradation and destruction of ecosystems. In addition to its ecological impact it has enormous potential to improve population health, socioeconomic well‐being, and the integrity of diverse national and ethnic cultures. In recognition of the critical role of restoration in ecosystem health, the United Nations (UN) declared 2021–2030 as the Decade on Ecosystem Restoration. We propose six practical strategies to strengthen the effectiveness and amplify the work of ecological restoration to meet the aspirations of the Decade: (1) incorporate holistic actions, including working at effective scale; (2) include traditional ecological knowledge (TEK); (3) collaborate with allied movements and organizations; (4) advance and apply soil microbiome science and technology; (5) provide training and capacity‐building opportunities for communities and practitioners; and (6) study and show the relationships between ecosystem health and human health. We offer these in the hope of identifying possible leverage points and pathways for collaborative action among interdisciplinary groups already committed to act and support the UN Decade on Ecosystem Restoration. Collectively, these six strategies work synergistically to improve human health and also the health of the ecosystems on which we all depend, and can be the basis for a global restorative culture.     

Causes of reintroduction failure of the brown treecreeper: Implications for ecosystem restoration

Reintroductions are conducted to re‐establish a self‐sustaining population of a species and contribute to ecosystem restoration. The brown treecreeper (Climacteris picumnus) reintroduction into two nature reserves in the Australian Capital Territory in south‐eastern Australia failed to meet its predetermined criteria for success. This occurred despite prior habitat restoration within the reserves where reintroduction occurred. Low survival of reintroduced brown treecreepers, particularly due to predation by native predators, has previously been highlighted as a key factor in the failure of the programme. We compared bird behaviour and habitat characteristics between the reintroduction reserves and the sites where brown treecreepers were sourced (which support stable brown treecreeper populations). We did not identify an indication of significantly higher predation pressure in the reintroduction reserves in comparison with the source sites. However, our results revealed that reintroduced individuals may be more vulnerable to predation because of an increased flight time to reach a refuge area. This was a result of a significantly lower number of refuge areas in logs and trees and a higher number of shrubs (which may obstruct escape paths and hinder detection of predators) in the reintroduction reserves compared with the source sites. We identified a lower ground foraging habitat quality in the reintroduction reserves because of lower numbers of ant mounds and lower areas of forageable ground. However, brown treecreepers were able to disperse extensively throughout the reserves and settle in areas with generally higher‐quality foraging habitat. Therefore, the negative effect of low ground foraging habitat quality would have been most pronounced immediately after release. This study emphasizes the inherent complexities of species reintroductions and ecosystem restoration. Despite experimental restoration activities within the reintroduction reserves, there were still deficiencies in habitat quality. We emphasize that further habitat restoration is required within these reserves to achieve more complete restoration.     

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.     

Restoration of California Native Grasses and Clovers: The Roles of Clipping,Broadleaf Herbicide,and Native Grass Density

One of the major challenges confronting grassland restoration of highly invaded communities is increasing the diversity of native species. There is surprisingly little research investigating how reconstructed native grasslands respond to common management techniques and how these techniques influence the relative establishment of both native grasses and forbs. Despite the diversity and wide distribution of native clovers in California, few practitioners incorporate them into grassland restoration plans. Conversely, non‐native clovers have been seeded extensively onto California rangelands. This study addresses the following questions: (1) Using readily available management tools, is there a strategy that can benefit the growth of both planted native bunchgrasses and seeded clovers? (2) Do native bunchgrasses compete with establishing clovers and non‐native grasses? (3) Do native and non‐native clovers differ in their response to management treatments or in their productivity? Plots were established to test three factors in different combinations over 3 years: (1) early spring clipping, (2) initial broadleaf herbicide, and (3) native bunchgrass planting density. Native and non‐native clovers were seeded in years 2 and 3. Early spring clipping did not have a significant effect on native bunchgrass cover, yet it did result in greater growth of native and non‐native clovers. The direction of the response to broadleaf herbicide changed between years for native bunchgrasses and was consistently negative for native clovers. Plots with higher native grass densities did not adversely affect the seeded clovers, yet non‐native grass cover was reduced. Native and non‐native clovers exhibited similar responses to clipping and established at similar densities.     

Restoration for variability: emergence of the habitat diversity paradigm in terrestrial ecosystem restoration

Ecosystem restoration implies focusing on multiple trophic levels and ecosystem functioning, yet higher trophic levels, that is, animals, are less frequently targeted by restoration than plants. Habitat diversity, the spatial heterogeneity between and within habitat patches in a landscape, is a well‐known driver of species diversity, and offers possible ways to increase species diversity at multiple trophic levels. We argue that habitat diversity is central in whole‐ecosystem restoration as we review its importance, provide a practical definition for its components, and propose ways to target it in restoration. Restoration targeting habitat diversity is used commonly in aquatic ecosystems, mostly to increase the physical diversity of habitats, meant to provide more niches available to a higher number of animal species. To facilitate the uptake of habitat diversity in terrestrial ecosystem restoration, we distinguish between compositional and structural habitat diversity, because different animal groups will respond to different aspects of habitat diversity. We also propose four methods to increase habitat diversity: varying the starting conditions to obtain divergent successional pathways, emulating natural disturbances, establishing keystone structures, and applying ecosystem engineer species. We provide two case studies to illustrate how these components and methods can be incorporated in restoration. We conclude that targeting habitat diversity is a promising way to restore habitats for a multitude of species of animals and plants, and that it should become mainstream in restoration ecology and practice. We encourage the restoration community to consider compositional and structural habitat diversity and to specifically target habitat diversity in ecosystem restoration.     

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.     

Harvester ant seed removal in an invaded sagebrush ecosystem: Implications for restoration

A better understanding of seed movement in plant community dynamics is needed, especially in light of disturbance‐driven changes and investments into restoring degraded plant communities. A primary agent of change within the sagebrush‐steppe is wildfire and invasion by non‐native forbs and grasses, primarily cheatgrass (Bromus tectorum). Our objectives were to quantify seed removal and evaluate ecological factors influencing seed removal within degraded sagebrush‐steppe by granivorous Owyhee harvester ants (Pogonomyrmex salinus Olsen). In 2014, we sampled 76 harvester ant nests across 11 plots spanning a gradient of cheatgrass invasion (40%–91% cover) in southwestern Idaho, United States. We presented seeds from four plant species commonly used in postfire restoration at 1.5 and 3.0 m from each nest to quantify seed removal. We evaluated seed selection for presented species, monthly removal, and whether biotic and abiotic factors (e.g., distance to nearest nest, temperature) influenced seed removal. Our top model indicated seed removal was positively correlated with nest height, an indicator of colony size. Distance to seeds and cheatgrass canopy cover reduced seed removal, likely due to increased search and handling time. Harvester ants were selective, removing Indian ricegrass (Achnatherum hymenoides) more than any other species presented. We suspect this was due to ease of seed handling and low weight variability. Nest density influenced monthly seed removal, as we estimated monthly removal of 1,890 seeds for 0.25 ha plots with 1 nest and 29,850 seeds for plots with 15 nests. Applying monthly seed removal to historical restoration treatments across the western United States showed harvester ants can greatly reduce seed availability at degraded sagebrush sites; for instance, fourwing saltbush (Atriplex canescens) seeds could be removed in <2 months. Collectively, these results shed light on seed removal by harvester ants and emphasize their potential influence on postfire restoration within invaded sagebrush communities.