International principles and standards for native seeds in ecological restoration

The growing demand for native seeds in ecological restoration and rehabilitation, whether for mining, forest, or ecosystem restoration, has resulted in a major global industry in the sourcing, supply, and sale of native seeds. However, there are no international guidance documents for ensuring that native seeds have the same standards of quality assurance that are regular practice in the crop and horticultural industries. Using the International Principles and Standards for the Practice of Ecological Restoration as a foundation document, we provide for the first time a synthesis of general practices in the native seed supply chain to derive the Principles and Standards for Native Seeds in Ecological Restoration (“Standards”). These practices and the underpinning science provide the basis for developing quality measures and guidance statements that are adaptable at the local, biome, or national scale. Importantly, these Standards define what is considered native seed in ecological restoration and highlight the differences between native seeds versus seeds of improved genetics. Seed testing approaches are provided within a logical framework that outline the many different dormancy states in native seed that can confound restoration outcomes. A “pro‐forma” template for a production label is included as a practical tool that can be customized for local needs and to standardize reporting to end‐users on the level of seed quality and germinability to be expected in a native seed batch. These Standards are not intended to be mandatory; however, the guidance statements provide the foundation upon which regulatory approaches can be developed by constituencies and jurisdictions.     

Seed storage: maintaining seed viability and vigor for restoration use

Effective seed storage after sourcing (harvesting or purchasing) is critical to restoration practitioners and native seed producers, as it is key to maintaining seed viability. Inadequate seed storage can lead to a waste of both natural and economic resources when seeds of poor quality are sown. When working with native species with unknown storage behavior, general assumptions can be made based on studies on related species, and standard practices may be applied with caution; however, an investigation should be conducted to understand if specific storage requirements are needed and for how long seeds can be stored before they lose significant viability. In this paper of the Special Issue Standards for Native Seeds in Ecological Restoration, we provide an overview of the key concepts in seed storage and the steps to take for effective storage of native seeds for restoration use.     

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.     

Lack of adequate seed supply is a major bottleneck for effective ecosystem restoration in Chile: friendly amendment to Bannister et al. (2018)

We argue that the need for a quality seed supply chain is a major bottleneck for the restoration of Chile's native ecosystems, thus supplementing the list of bottlenecks proposed by Bannister et al. in 2018. Specifically, there is a need for defining seed transfer zones, developing standards and capacities for properly collecting and storing seeds, reducing information gaps on seed physiology and longevity, and implementing an efficient seed supply chain with certification of seed origin and quality. Without such capacities, countries are unlikely to meet their restoration commitments. Although we focus on bottlenecks in Chile, the issues we raise are relevant to other countries and thus the global agenda for ecological restoration.     

How policies constrain native seed supply for restoration in Brazil

Large‐scale ecological restoration programs across the world involve a voluminous demand for native seeds of diverse native plant species. In this article, we explore how institutional systems have operated and impacted native seed supply in Brazil. Native seed supply for restoration is essentially a community‐based activity which faces broad barriers to operating within regulations because of requirements for excessive and costly technical documentation, scarcity of seed laboratories, and lack of instructions for native seed quality testing. Although decentralized seed networks have stimulated arrangements for local organizations to promote seed supply, policies constrain the development of local capacities and the ongoing sustainability of these organizations. These conditions have resulted in a vast network of informal collectors and producers who are largely “invisible” and unknown to the regulatory authorities. Policies have decentralized responsibilities from the state without devolving decision‐making power to the multiple stakeholders engaged in policy elaboration. The policies maintain the centralized regulation of native seed supply. After examining Brazilian seed networks' experiences and conducting discussions with stakeholders and experts, we suggest adapting the current regulations to more local level contexts, encompassing the following strategies: (1) ensuring native seed origin and identity; (2) relaxation of the laboratory accreditation process for native seed quality assurance; (3) fostering seed markets for restoration; (4) research to provide technological innovation; (5) supporting local, diverse, and small seed‐based businesses.     

Ensuring seed quality in ecological restoration: native seed cleaning and testing

Seeds are a critical and limited resource for restoring biodiversity and ecological function to degraded and fragmented ecosystems. Cleaning and quality testing are two key steps in the native seed supply chain. Optimizing the practices used in these steps can ensure seed quality. Post‐collection handling of seeds can have a profound impact on their viability, longevity in storage, and establishment potential. The first section of this article describes seed cleaning, outlines key considerations, and details traditional and novel approaches. Despite the growth of the native seed industry and the need for seed quality standards, existing equipment and standards largely target agricultural, horticultural, and commercial forestry species. Native plant species typically have complex seed traits, making it difficult to directly transfer existing cleaning and quality standards to these species. Furthermore, in ecological restoration projects, where diversity is valued over uniformity crop standards can be unsuitable. We provide an overview and recommendations for seed quality testing (sampling, purity, viability, germinability, vigor), identity reporting, and seed transfer as well as highlight the need to implement internationally recognized standards for certification for native seeds. Novel and improved cleaning and testing methods are needed for native species from a range of ecosystems to meet the challenges and goals of the United Nations Decade on Ecosystem Restoration. The guidelines outlined in this article along with others in the Special Issue of Restoration Ecology “Standards for Native Seeds in Ecological Restoration” can serve as a foundation for this critical work.     

Collection and production of native seeds for ecological restoration

The global push to achieve ecosystem restoration targets has resulted in an increased demand for native seeds that current production systems are not able to fulfill. In many countries, seeds used in ecological restoration are often sourced from natural populations. Though providing seed that is reflective of the genetic diversity of a species, wild harvesting often cannot meet the demands for large‐scale restoration and may also result in depletion of native seed resources through over harvesting. To improve seed production and decrease seed costs, seed production systems have been established in several countries to generate native seeds based on agricultural or horticultural production methods or by managing natural populations. However, there is a need to expand these production systems which have a primary focus on herbaceous species to also include slower maturing shrub and tree seed. Here we propose that to reduce the threat of overharvest on the viability of natural populations, seed collection from natural populations should be replaced or supplemented by seed production systems. This overview of seed production systems demonstrates how to maximize production and minimize unintended selection bias so that native seed batches maintain genetic diversity and adaptability to underpin the success of ecological restoration programs.     

Spicing up restoration: can chili peppers improve restoration seeding by reducing seed predation?

Seed predation by rodents presents a significant barrier to native plant recruitment and can impede restoration seeding efforts. In nature, some plants contain secondary defense compounds that deter seed predators. If these natural defense compounds can be applied to unprotected seeds to inhibit rodent granivores, this approach could improve restoration seeding. Capsaicin is the active ingredient in chili pepper (Capsicum spp.) seeds that creates the burning sensation associated with human consumption of hot peppers. This compound has a similar effect on other mammals and is believed to have evolved as a deterrent to rodent seed predators. We used seed‐coating techniques to attach powder ground from Bhut Jolokia (Capsicum chinense) peppers to native plant seeds and evaluated the efficacy of these seed coatings for deterring rodent seed predation and enhancing native plant recruitment using laboratory and field experiments. Laboratory feeding trials demonstrated that native deer mice (Peromyscus maniculatus) consumed far fewer pepper‐coated seeds compared to untreated control seeds. Field seed‐addition experiments consistently demonstrated that rodent seed predation reduced native plant recruitment over the 4‐year study. Coating techniques used in the first 3 years were not persistent enough to reduce rodent seed predation effects on plant recruitment. However, a more persistent coating applied in conjunction with late‐winter sowing negated rodent seed predation effects on recruitment in year 4. Our results demonstrate that coating seeds with natural plant defense compounds may provide an effective, economical way to improve the efficacy of plant restoration by deterring seed predation by ubiquitous rodent granivores.     

Ten years of native seed certification in Germany – a summary

Many renaturation projects and compensation areas are based on the use of seeds from regional indigenous wild plants; in the following: native or regional seeds. Despite this, such seeds make up only a small proportion of the total number of seeds used for greening projects; in Germany, for example, it is only around 1% (=200 t per year). Although the market for regional seeds is small, it is highly competitive. High‐priced native seeds compete with flower mixes of unspecified origin and can only be differentiated from them by reliable quality seals. A quality assurance system based on seed legislation (EU Directive 2010/60, preservation mixtures) has been developed in a few European countries. However, quality assurance ends with the sale of the seeds. Thus, seed use remains unmonitored, and often unsuitable material, or material foreign to the region, is planted in restoration areas. Unfortunately, nature conservation has not made seed‐based restoration one of its key issues, neither at the European nor at the national level. Currently there are many different local and regional standards, methods and private certificates that are confusing for users and which provide little continuity and predictability for producers. We recommend the establishment of an EU directive or a broadly agreed recommendation to the EU member states, spearheaded by nature conservation, which would define the standards for producing and using native seeds (e.g. harmonised regions that cross national borders, quality regulations). At the same time, wild plant interest groups should combine existing structures in order to strengthen seed‐based restoration through international cooperation.     

Seed enhancement: getting seeds restoration‐ready

Seed enhancement technologies such as seed priming and seed coating, developed by the agricultural seed industry, are standard procedures for the majority of crop and horticultural seeds. However, such technologies are only just being evaluated for native plant seeds despite the potential benefits of such treatments for improving restoration effectiveness. Key approaches applicable to native seed include: (1) seed priming, where seeds are hydrated under controlled conditions, and (2) seed coating, in which external materials and compounds are applied onto seeds through a diversity of treatments. These technologies are commonly employed to accelerate and synchronize germination and to improve seed vigor, seedling emergence, establishment, and to facilitate mechanized seed delivery to site, through standardizing seed size and shape. Seed enhancement technologies have now been tested on native seeds to overcome logistical and ecological barriers in restoration. However, further research is needed to extend the application of seed enhancements to a broader array of species, ecosystems, and regions as well as to evaluate new and innovative approaches such as the incorporation of beneficial soil microorganisms and plant growth regulators in the coatings. As techniques in native seed enhancement develop, these approaches need to be capable of being scaled‐up to provide the tonnages of seed required for global restoration.     

Seed planning,sourcing, and procurement

Ensuring the availability of adequate seed supplies of species and sources appropriate for restoration projects and programs necessitates extensive science‐based planning. The selection of target species requires a review of disturbance conditions and reference areas, development of a reference model, and consideration of specific objectives, timeframes, available resources, and budgets as well as the performance of prospective species in past restoration efforts. Identification of seed sources adapted to site conditions is critical to provide for short‐term establishment and long‐term sustainability. Seed zones and plant movement guidelines provide tools for sourcing plant materials with reduced risk of maladaptation. A seed zone framework also facilitates seed use planning and contributes to stability and predictability of the commercial market, thereby reducing costs and improving the availability of adapted seed supplies. Calculating the amount of seed required for each species is based on seed quality (viability, purity), seed weight, expected seedling establishment, and desired composition of the seeding. If adequate collections from wildland stands are not feasible, then seed increase in seed fields or use of nursery stock may be warranted. Adherence to seed collection and seed production protocols for conserving genetic diversity is critical to protect genetic resources and buffer new seedings and plantings against environmental stressors. Maintenance of genetic diversity becomes even more critical considering current or expected climate change impacts. Collaboration and partnerships can benefit seed selection and procurement programs through sharing of information, coordination in project planning, and increasing the availability of native seed.     

Comparison of seed and seedling functional traits in native Helianthus species and the crop H. annuus (sunflower)

• Seed functional traits of native Helianthus species contribute towards ecosystem services but limitations to their use in managed programmes exist. Many perennial Helianthus possess seed dormancy. The ability for germination to occur under different temperature and drought conditions, as well as the capacity of germinated seeds to convert into normal seedlings is rarely considered. Our aim was to identify and quantify these constraints through functional trait analyses.
• In five seed lots of native Helianthus (four perennial and one annual) and five genotypes of sunflower (H. annuus) for comparison, dormancy, thermal and hydro thresholds and times, morphology, mass, oil content and conversion into normal seedlings were quantified. The influence of the seed collection site environment on these traits was also explored.
• Seed dormancy of the perennial species was overcome by scarification followed by germination in 5 mm GA₃. Thermal and hydro‐time analyses revealed slower germination for the native seed lots (>1350 °Ch) in comparison to the sunflower genotypes (<829.9 °Ch). However, native seed lots had a higher capacity to convert into normal seedlings at high temperatures and low water potentials than sunflower genotypes. For the native seed lots, the average monthly temperature of the collection site was negatively correlated with thermal time.
• Variability in seed functional traits of native Helianthus and greater capacity for germinated seeds to convert into normal seedlings suggests they are better equipped to cope with high temperature and drought scenarios than sunflower. Effective dormancy alleviation is required to facilitate the use of native Helianthus species.

     

Non-native mammals are weak candidates to substitute ecological function of native avian seed dispersers in an island ecosystem

Although prominent examples exist of non-native species causing substantial ecological harm, many have neutral or positive effects, including filling surrogate roles once performed by extinct native organisms. We tested the ecological roles of two non-native mammals as seed dispersers or seed predators in Guåhan, which, due to invasive brown tree snakes (Boiga irregularis), is devoid of native seed dispersers–birds and bats. We conducted feeding trials with captive rats (Rattus spp.), which are present but uncommon due to predation by snakes, and pigs (Sus scrofa), which are abundant. We examined if and how they interacted with common forest fruits. We then compared how any gut-passed or animal-handled seeds germinated compared to seeds left in whole fruit or depulped seeds. Rats and pigs interacted with most of the fruits and seeds (>80%) that they were fed. Of those, most seeds were destroyed—78% for rats and 90% for pigs, across both native and non-native plant species. Compared to seeds germinating within whole fruits, rats improved germination of the seeds that they handled without ingesting, while pigs diminished the germination of seeds that they handled. The small percentage of seeds (approximately 1.5% for rats and 5% for pigs) that survived gut passage germinated in higher proportions than those in whole fruits. Percentages of seed survival to germination are lower than found in similar studies with native avian frugivores. Our results indicate that pigs and rats have mixed effects on seeds, but are not suitable surrogates for native seed dispersers.     

Native seed trade of herbaceous species for restoration: a European policy perspective with global implications

With the need to meet ambitious restoration targets, an improved native seed sector for the production of herbaceous species with a practical and supportive policy framework is recognized. We evaluated the current “ready‐made” policy frameworks in Europe regarding the native seed supply of herbaceous species and found them to be, generally, unsatisfactory for both producers and users. Initially, such policies were designed for fodder seed and relate to distinctness, uniformity, and stability, traits that do not reflect the genetic heterogeneity of native species required for ecological restoration. Until recently, more suitable certification standards were designed to multiply fodder seed for preservation of the natural environment; however, due to the disparateness of the seed market in Europe, this policy is rarely practical and fails to encompass all herbaceous native species often resulting in unregulated seed sales. We recommend a new or adapted native seed policy constructed through a participatory or bottom‐up approach and supported through the formation of widely based trade associations. Such a policy could stimulate the native seed trade with concomitant impacts on the speed of improving ecosystem services.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

Woody structure facilitates invasion of woody plants by providing perches for birds

Woody encroachment threatens prairie ecosystems globally, and thus understanding the mechanisms that facilitate woody encroachment is of critical importance. Coastal tallgrass prairies along the Gulf Coast of the US are currently threatened by the spread of several species of woody plants. We studied a coastal tallgrass prairie in Texas, USA, to determine if existing woody structure increased the supply of seeds from woody plants via dispersal by birds. Specifically, we determined if (i) more seedlings of an invasive tree (Tridacia sebifera) are present surrounding a native woody plant (Myrica cerifera); (ii) wooden perches increase the quantity of seeds dispersed to a grassland; and (iii) perches alter the composition of the seed rain seasonally in prairie habitats with differing amounts of native and invasive woody vegetation, both underneath and away from artificial wooden perches. More T. sebifera seedlings were found within M. cerifera patches than in graminoid‐dominated areas. Although perches did not affect the total number of seeds, perches changed the composition of seed rain to be less dominated by grasses and forbs. Specifically, 20–30 times as many seeds of two invasive species of woody plants were found underneath perches independent of background vegetation, especially during months when seed rain was highest. These results suggest that existing woody structure in a grassland can promote further woody encroachment by enhancing seed dispersal by birds. This finding argues for management to reduce woody plant abundance before exotic plants set seeds and argues against the use of artificial perches as a restoration technique in grasslands threatened by woody species.     

Seed Removal and Predation as Factors Affecting Seed Availability of Tree Species in Degraded Habitats and Restoration Plantings in Rainforest Areas of Queensland,Australia

The post‐dispersal removal or predation of seeds of native tree species was investigated in Queensland, Australia, at degraded habitats and rainforest restoration sites where direct seeding might be used to facilitate tree regeneration (old fields or open habitats, lantana thicket, rainforest edge, and 5‐ and 10‐year‐old restoration plantings). Seed removal/predation was assessed in relation to tree seed weight and canopy density of the habitats during the wet season period. Results indicated that seed removal/predation imposes limitations on seed availability, particularly for small seeded species. In most situations, larger seeds were less removed/predated, most likely due to the more limited range of large seed consumers. The use of large, hard‐coated seeds may potentially reduce seed loss in open situations (from both seed removal and desiccation), unless large seed consumers frequent the site. Canopy cover exerted an influence on seed removal/predation, though trends varied in relation to site and the time of season. Broadcast sowing of seed under planted tree canopies at the more advanced stages of closure may in some areas result in higher seed removal/predation. Likewise, seeding in areas dominated by woody weeds may result in high seed losses to consumers such as rodents. Results suggested that undertaking direct seeding to coincide with the maximal period of fruit production may in some situations be beneficial to minimize seed loss. Overall, site context, canopy cover, and species selection appear to be important considerations when aiming to reduce loss of seeds to animal seed consumers in restoration work.     

Message in a bottle: Inadvertent loss of seeds of native grassland species as a result of rudimentary long‐term storage

Many practitioners are likely to have collected seeds with the intention of using that seed for conservation and/or restoration plantings, but have not got around to using the seed, sometimes for many years. Currently, it is not clear what species have short‐lived or long‐lived seed when stored under rudimentary conditions such as in paper bags or in a refrigerator. We report the germinability of 12 temperate native grassland species, comprising 16 populations, whose seeds were collected with the purpose of raising seedlings to plant into the wild, but whose seeds were subsequently stored at ~2–4°C for 25+ years. We conclude that most of the grassland species that we assessed do not have viable seed after 25 years when stored in such conditions; only two species germinated despite evidence that seed germinates well for most species when first collected. Inadvertent loss of seeds of as a result of long‐term storage is most likely in readily germinable species (e.g. members of the Asteraceae). The ways in which seeds are stored by practitioners deserves consideration given the risk of seed mortality with long‐term storage under rudimentary conditions.     

Review of seed pelletizing strategies for arid land restoration

Arid lands face numerous restoration challenges due to infrequent and irregular precipitation which impacts plant germination, growth, and survival. Abundant seed predators, harsh surface conditions, and native seeds often poorly suited to mechanical distribution compound restoration challenges. Seed pellets (a.k.a. pods, seed balls, and seed bombs), an aggregation of clay, soil, water, and multiple seeds, have the potential to reduce some of the challenges. However, no formal guidelines based on an aggregated research review exist. Available publications, both peer‐reviewed and any public domain, were reviewed. Of the 24 publications found (some describing multiple tests), seed pellets tested on rangelands had 9 negative effects, 10 neutral effects, and 8 positive effects. Greenhouse testing showed five negative effects, two neutral effects, and four positive effects. Forest systems had three neutral effects and two positive effects. Advantages in mechanical distribution were not well quantified and other effects were lumped together in germination and seedling establishment totals. The use of pelleted seed would benefit from a more mechanistic understanding of the interaction between the pellets and the systemic constraints. As long as the pellets themselves do not greatly reduce germination or establishment, they appear a potentially viable method for improving seed distribution and seeding efficiency, protecting seeds, and adding amendments. While the evidence is far from conclusive, seed pellets may also hold some advantages in increasing germination and establishment.     

Avian Seed Dispersal of an Invasive Shrub

The incorporation of an animal-dispersed exotic plant species into the diet of native frugivores can be an important step to that species becoming invasive. We investigated bird dispersal of Lonicera maackii, an Asian shrub invasive in eastern North America. We (i) determined which species of birds disperse viable L. maackii seeds, (ii) tested the effect of gut passage on L. maackii seeds, and (iii) projected the seed shadow based on habitat use by a major disperser. We found that four native and one exotic bird species dispersed viable L. maackii seeds. Gut passage through American robins did not inhibit germination, but gut passage through cedar waxwings did. American robins moved mostly along woodlot edges and fencerows, leading us to project that most viable seeds would be defecated in such habitats, which are very suitable for L. maackii. We conclude that L. maackii has been successfully incorporated into the diets of native and exotic birds and that American robins preferentially disperse seeds to suitable habitat.