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.     

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.     

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.     

Community‐based native seed production for restoration in Brazil – the role of science and policy

• Large‐scale restoration programmes in the tropics require large volumes of high quality, genetically diverse and locally adapted seeds from a large number of species. However, scarcity of native seeds is a critical restriction to achieve restoration targets.
• In this paper, we analyse three successful community‐based networks that supply native seeds and seedlings for Brazilian Amazon and Cerrado restoration projects. In addition, we propose directions to promote local participation, legal, technical and commercialisation issues for up‐scaling the market of native seeds for restoration with high quality and social justice.
• We argue that effective community‐based restoration arrangements should follow some principles: (i) seed production must be based on real market demand; (ii) non‐governmental and governmental organisations have a key role in supporting local organisation, legal requirements and selling processes; (iii) local ecological knowledge and labour should be valued, enabling local communities to promote large‐scale seed production; (iv) applied research can help develop appropriate techniques and solve technical issues.
• The case studies from Brazil and principles presented here can be useful for the up‐scaling restoration ecology efforts in many other parts of the world and especially in tropical countries where improving rural community income is a strategy for biodiversity conservation and restoration.

     

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.     

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.     

Optimising seed processing techniques to improve germination and sowability of native grasses for ecological restoration

• Grasslands across the globe are undergoing expansive degradation due to human impacts and climate change. If restoration of degraded native grassland is to be achieved at the scale now required, cost‐effective means for seed‐based establishment of grass species is crucial. However, grass seeds present numerous challenges associated with handling and germination performance that must be overcome to improve the efficiency of seeding. Previous research has demonstrated that complete removal of the palea and lemma (husk) maximises germination performance, hence we investigated the effects of complete husk removal on seed handling and germination of four temperate Australian grass species.
• Three techniques were tested to remove the husk – manual cleaning, flaming or acid digestion (the latter two followed by a manual cleaning step); these techniques were refined and adapted to the selected species, and germination responses were compared.
• The complete removal of the husk improved seed handling and sowability for all species. Germination was improved in Microlaena stipoides by 19% and in Rytidosperma geniculatum by 11%. Of the husk removal methods tested, flaming was detrimental to seed germination and fatal for one species (R. geniculatum). Compared to manual cleaning, sulphuric acid improved the overall efficacy of the cleaning procedure and increased germination speed (T50) in Austrostipa scabra, Chloris truncata and M. stipoides, and improved final germination in R. geniculatum by 13%.
• The seed processing methods developed and tested in the present study can be applied to grass species that present similar handling and germination performance impediments. These and other technological developments (seed coating and precision sowing) will facilitate more efficient grassland restoration at large scale.

     

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.     

亏缺灌溉对干旱区两种乡土植物种子生产性能及其水分利用效率的影响

科学灌溉对植物种子生产具有重要意义。本研究以荒漠草原优良乡土植物沙芦草和牛枝子为对象,以充分灌溉为对照,探究不同生育时期亏缺灌溉对两种牧草种子生产和水分利用效率的影响。结果表明: 与对照相比,亏缺灌溉下两种植物土壤含水率下降,其中沙芦草土壤含水率下降主要发生在0～60 cm土层,牛枝子土壤水分下降未出现明显的分层现象。亏缺灌溉下沙芦草种子产量各构成因子差异均显著,开花期亏缺灌溉种子产量最高;牛枝子仅生殖枝数、小花数和荚果数差异显著,种子产量各处理差异不显著。相关分析显示,沙芦草种子产量与生殖枝数(r=0.776)、小穗数(r=0.717)呈显著正相关;牛枝子花序数与生殖枝数呈极显著负相关(r=-0.685),与小花数呈显著正相关(r=0.412)。与充分灌溉相比,亏缺灌溉下两种乡土植物种子生产耗水量减少,水分利用效率提高,其中,沙芦草开花期亏缺灌溉水分利用效率提高最多(32.9%);牛枝子分枝期亏缺灌溉提高最多(27.4%)。因此,适当亏缺灌溉可以提高两种植物水分利用效率。从水分利用效率和种子产量来看,干旱区沙芦草和牛枝子种子人工繁育时可采取亏缺灌溉,适宜亏缺的生育期分别为开花期和分枝期。     

采用本土蔬菜种子替代水堇评价污泥有机肥腐熟度

旨在筛选出对城市污泥毒性物质敏感、与国际通用的水堇种子相似、可用于评价处理后的城市污泥有机肥的植物毒性的本土植物种子。试验采用我国常见的5个科的17种蔬菜种子,检测了不同腐熟度的城市污泥堆肥以及经植物处理的城市污泥产品的种子发芽系数,同时以水堇(Lepidium sativum L.)种子发芽系数为标准作比较分析。结果表明,对于未腐熟的污泥堆肥,油青四九菜心(Brassica parachinensis Bailey)、青江白菜(Brassica campestris Pekinensis)、京丰一号甘蓝(Brassica oleracea L.)等种子对污泥植物毒性较敏感,与水堇种子较相似。对于植物处理后的污泥产品,青江白菜和京丰一号甘蓝与水堇种子又有明显差异,只有油青四九菜心与进口水堇种子对污泥腐熟度的响应未表现出明显差异。而且油青四九菜心种子发芽系数与水堇之间的直线相关系数R²达到0.7465,水堇种子发芽系数0.5时对应于的该菜心种子发芽系数为0.53,因此较适合代替目前广泛用于检测污泥有机肥腐熟度的进口水堇种子。     

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.     

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.     

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.     

Rapid evolution in native plants cultivated for ecological restoration: not a general pattern

• The growing number of restoration projects worldwide increases the demand for seed material of native species. To meet this demand, seeds are often produced through large‐scale cultivation on specialised farms, using wild‐collected seeds as the original sources. However, during cultivation, plants experience novel environmental conditions compared to those in natural populations, and there is a danger that the plants in cultivation are subject to unintended selection and lose their adaptation to natural habitats. Although the propagation methods are usually designed to maintain as much natural genetic diversity as possible, the effectiveness of these measures have never been tested.
• We obtained seed of five common grassland species from one of the largest native seed producers in Germany. For each species, the seeds were from multiple generations of seed production. We used AFLP markers and a common garden experiment to test for genetic and phenotypic changes during cultivation of these plants.
• The molecular markers detected significant evolutionary changes in three out of the five species and we found significant phenotypic changes in two species. The only species that showed substantial genetic and phenotypic changes was the short‐lived and predominantly selfing Medicago lupulina, while in the other, mostly perennial and outcrossing species, the observed changes were mostly minor.
• Agricultural propagation of native seed material for restoration can cause evolutionary changes, at least in some species. We recommend caution, particularly in selfing and short‐lived species, where evolution may be more rapid and effects may thus be more severe.

     

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.     

Soil seed bank in different vegetation types in the Loess Plateau region and its role in vegetation restoration

In the Loess Plateau region, soil erosion is a serious problem. Vegetation restoration is an effective approach to control soil erosion and improve ecosystems. The soil seed bank generally plays an important role in vegetation restoration after disturbance. Thus, we reviewed soil seed bank studies to reveal the soil seed bank characteristics and its role in vegetation restoration in three vegetation types (forest, forest‐steppe, and steppe). We selected 38 seed bank studies and analyzed several seed bank characteristics, such as seed density, species composition, and the relationship between seed size and seed bank. We also assessed the role of the soil seed bank in vegetation restoration. The soil seed bank density ranged from 2,331 ± 1,993 to 6,985 ± 4,047 seeds/m² among the different vegetation types. In the soil seed bank, perennial herbs and grasses accounted for 51.5% of the total species. Native species that were dominant or common in the standing vegetation usually had relatively high seed bank densities. Moreover, species with smaller seeds generally had higher soil seed bank densities. The present study indicates that the soil seed bank plays a significant role in spontaneous vegetation restoration, especially during the early successional stages in abandoned slope farmlands and grazing‐excluded grasslands. However, species with large seeds or transient soil seed banks should be reintroduced through seeding to accelerate target species restoration. More studies on soil seed banks need to be conducted to comprehensively reveal their characteristics.     

Hydrothermal thresholds for seed germination in winter annual forbs from old‐field Mediterranean landscapes

• Under Mediterranean climates with dry‐hot summers and cool‐wet winters, many forbs with potential for habitat restoration are winter annuals, but there is little information about their germination.
• We performed laboratory germination experiments on 13 ruderal dicots native to Andalusia (southern Spain). We measured the germination of recently harvested seeds from natural populations across nine temperature treatments (from 5 to 35 °C, constant and alternate); two storage periods; and eight water stress treatments (from 0 to ?1.0 MPa). We then calculated the hydrothermal thresholds for seed germination.
• Final germination ranged from 0–100% and results were mixed in response to temperature. Base temperature was below 6 °C, optimal temperature was around 14 °C and the ceiling temperature around 23 °C. For five species, 10 months of storage improved total germination, indicating a dormancy‐breaking effect, but the other species did not respond or had their germination reduced. All species were relatively tolerant to water stress, with base water potential ranging from ?0.8 to ?1.8 MPa.
• Our results suggest that hydrothermal germination thresholds, rather than physiological dormancy, are the main drivers of germination phenology in annual forbs from Mediterranean semi‐dry environments. The variation in germination responses of these forb species differs from winter annual grasses, but their seeds are all suitable for being stored before restoration.

     

Variations in seed germination behavior of Phleum hirsutum subsp. ambiguum and possible applications in semi-natural grassland restoration

Phleum hirsutum Honck. subsp. ambiguum (Ten.) Tzvelev is an important component of the vegetation of some semi-natural grassland habitats of central and southern Italy and especially of those referring to the 6210^* habitat (Directive 92/43/EEC). Seeds of this species are required for the execution of restoration works concerning this habitat. Therefore, the recovery and the study of autochthonous germplasm are needed. This study focused on the intra-specific variations in seed mass and size and in seed germination behavior in this species since the knowledge of these aspects can be very useful in the planning of restoration projects. The present research evidenced significant differences in mean seed mass and size in the populations studied that were correlated to the precipitation values of their collection sites. Studied seeds are able to germinate under a wide range of temperatures. They reached their maximum germination between 5 and 25°C and are capable of germination even at 30°C but with a remarkable reduction in germination percentage. Temperature was found to strongly influence germination, especially in terms of the speed. Light was found to enhance germination in almost all the populations studied while the removal of outer covering structures was found to be of no effect.     

Dormancy and germination: making every seed count in restoration

From 50 to 90% of wild plant species worldwide produce seeds that are dormant upon maturity, with specific dormancy traits driven by species' occurrence geography, growth form, and genetic factors. While dormancy is a beneficial adaptation for intact natural systems, it can limit plant recruitment in restoration scenarios because seeds may take several seasons to lose dormancy and consequently show low or erratic germination. During this time, seed predation, weed competition, soil erosion, and seed viability loss can lead to plant re‐establishment failure. Understanding and considering seed dormancy and germination traits in restoration planning are thus critical to ensuring effective seed management and seed use efficiency. There are five known dormancy classes (physiological, physical, combinational, morphological, and morphophysiological), each requiring specific cues to alleviate dormancy and enable germination. The dormancy status of a seed can be determined through a series of simple steps that account for initial seed quality and assess germination across a range of environmental conditions. In this article, we outline the steps of the dormancy classification process and the various corresponding methodologies for ex situ dormancy alleviation. We also highlight the importance of record‐keeping and reporting of seed accession information (e.g. geographic coordinates of the seed collection location, cleaning and quality information, storage conditions, and dormancy testing data) to ensure that these factors are adequately considered in restoration planning.     

The effect of micronutrient supply on the growth and seed production of guayule (Parthenium argentatum Gray)

The effect of micronutrient supply on the growth and seed production of guayule (Parthenium argentatum Gray) was investigated. A deficiency of boron, but no other trace element, significantly reduced the growth, seed production and the percentage of seeds that germinated. The optimum concentration of B for seed production was between 20 and 100 μM. Results indicate that guayule may benefit from B fertilisation.