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.     

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.     

Recovery of rain forest soil seed banks under different reforestation pathways in eastern Australia

Summary Seed availability is a major factor limiting the recruitment of rain forest to cleared land, but little is known about the composition of the soil seed bank under different reforestation pathways. We quantified changes in the viable soil seed bank following rain forest clearing and pasture establishment and subsequent reforestation in subtropical eastern Australia. Major reforestation pathways in the region include planting of a diverse suite of native trees for ecological restoration purposes, autogenic regrowth dominated by the non‐native tree Camphor Laurel (Cinnamomum camphora) and management of this regrowth to accelerate the development of a native tree community. These pathways differ considerably in cost: restoration plantings are expensive, autogenic regrowth is free, whilst managing regrowth generally costs much less than restoration plantings. We surveyed five sites within each of three reforestation pathways as well as reference sites in remnant rain forest and pasture. The composition of the seed bank was determined by germinating plants from soil samples collected from each site. Germinants were classified into several functional groups according to life form, origin, dispersal mode and successional stage. The majority of functional groups varied significantly in abundance or richness between rain forest and pasture sites. Most of the functional groups that varied between rain forest and pasture were restored to values similar to rain forest by at least one of the three reforestation pathways examined. The species richness of native woody plants in the soil seed bank was slightly higher in restoration plantings than in autogenic or managed regrowth; nevertheless, the species richness and abundance of native woody plants and vines were higher in the seed bank of autogenic regrowth than pasture, and both attributes were enhanced by the management of regrowth sites. The results of this study show that autogenic regrowth can make an important contribution to rain forest restoration at a landscape scale. The optimal reforestation approach or mix of approaches will depend on the desired rate of recovery and the resources available for restoration.     

Polycross Seed of Genetically Diverse Smooth Cordgrass (Spartina alterniflora) for Erosion Control and Habitat Restoration

Although seed‐based planting is common in crop systems, it is relatively a new concept in coastal erosion control and habitat restoration. This paper discusses the potential use of seed‐based revegetation to accommodate large‐scale erosion control using a highly diverse population of smooth cordgrass (Spartina alterniflora) from controlled polycross to attain desirable genetic diversity suitable for habitat restoration. Seed‐based restoration provides a more versatile alternative approach to the current clonal revegetation technique in many regions, including the Gulf Coast of the United States. The objectives of this study were to (1) describe seed production and cultural aspects of the polycross population and (2) discuss the potential use of polycross seeds for direct seeding applications and other innovative restoration approaches using seed‐derived planting materials. The polycross population was produced using 15 genetically diverse and high‐seed producing smooth cordgrass lines selected from native populations. The average seed set of the polycross population was 58.5 ± 6.3% with an average germination rate of 82.2 ± 9%. As comparison, Vermilion, the only available smooth cordgrass cultivar, has a seed set of 20.6 ± 5% and a germination rate of 35 ± 8%. The average yield of S. alterniflora seed from the polycross population was 277.5 kg/ha, which is equivalent to approximately 26 million viable seeds. Seed can be stored in 100% humidity at a temperature of 2 ± 1°C for 6–9 months.     

A Molecular and Fitness Evaluation of Commercially Available versus Locally Collected Blue Lupine Lupinus perennis L. Seeds for Use in Ecosystem Restoration Efforts

Dependence on wild seed sources is often impractical for large‐scale habitat restoration programs. Reliance on commercial seed supplies of unknown provenance and fitness is thereby warranted. Little consideration has been given, however, to how the large volumes of seed required should be sourced. We evaluated commercial and locally collected seed sources for potential use in a New York State‐based, landscape‐scale program for restoring blue lupine Lupinus perennis. Through analysis of microsatellite markers we determined that “native” lupine designations by some commercial suppliers were in fact interspecific hybrids and therefore unreliable; at least two commercial sources, however, were genetically as close to native New York populations as native New York populations were to one other. Common garden experiments revealed that seed source influenced first‐year overwintering survival and subsequent height growth of surviving plants; seed sources more closely related genetically to native New York populations survived better and produced more stems per individual in the field in the area targeted for restoration. We conclude that (1) commercial suppliers often but not always offer reliably characterized seed sources of sufficient genetic similarity to native populations to warrant their use in restoration projects and (2) genetic affinity of potential seed stock to native populations is positively related to its fitness in the environment targeted for restoration.     

Should we be more critical of remnant seed sources being used for revegetation?

Summary A challenge for land managers restoring degraded agricultural landscapes across southern Australia will be to ensure the viability of remnant vegetation while simultaneously supplying the quantities of appropriate seed required for revegetation. To ensure such revegetation programs have the best chance of success, seed that is both genetically diverse and locally adapted will be required. Identifying suitable seed sources can be particularly difficult in regions where local seed sources are restricted to small and isolated remnants. Gold‐dust Wattle (Acacia acinacea Lindl.) is a key revegetation species in the Deniliquin region of New South Wales; however, broadscale land clearing in the area has limited local seed sources to a few remnant stands. Field‐based experience suggests that revegetation success may depend upon the source of seed used, raising the question of whether differences in the germination and survival of seed reflect functional problems within these source populations. To test this possibility, seed was collected from 15 mothers in each of three seed sources regularly used for local restoration programs. Seed quality for each mother was assessed in terms of seed production and seedling fitness. In addition, genetic diversity and mating system parameters were determined to assess whether these explained the seed quality responses observed. Differences among the seed sources with respect to seed quality were generally congruent with field‐based predictions. High levels of correlated paternity in the two poorly performing seed sources probably reflect limited mating availability due to smaller population sizes and genetic incompatibility being mediated by a self‐incompatible reproductive strategy. Further research is now required to determine whether observed variability in the quality of seed from remnant vegetation in degraded landscapes is compromising revegetation efforts, and to help practitioners develop strategies to critically evaluate their seed sources.     

美国对入侵种的管理对策

外来种入侵是产生当前世界生物多样性危机的重要原因之一。目前外来种入侵问题已经引起人们的普遍关注。1999年2月美国联邦政府发布总统令,号召联邦政府机构执行人员防止外来入侵种所造成的严重环境威胁,指示有关职能部门行使职权阻止引入入侵种并且恢复本地种。总统令指示建立一个入侵物种理事会,并就理事会、联邦机构的职责做了相应界定。美国农业部等部门支持的入侵植物管理国家策略项目,提出了针对入侵种的三项国家目标一入侵种的预防、控制和本地种的恢复。国家管理策略的制定不同层次的人们提供了合作、教育和研究的机遇。美国政府部门以及有关社会团体已经就外来种入侵危害防范积极展开行动,包括制定标准防止入侵种进入,控制、根除入侵种,向民众进行有关外来种入侵的教育,增加财政年度预算等等。     

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.

     

喀斯特乡土树种伊桐的种实性状及幼苗生长特征

研究了喀斯特乡土树种伊桐的果实性状、种子性状、播种萌发和幼苗生长等的基本特征。结果表明:伊桐果实单果鲜重63.20～149.50g,其长度和宽度则分别为55.59～86.60mm和38.59～51.80mm,单果出种量19.95～57.15g,出籽率31.57%～40.81%,单果种子数343～690粒,单粒种子均重0.0582～0.0857g,其出籽率、单果种子数和单粒种子均重与果实重量存在一定程度的正相关;伊桐种子的长、宽平均分别为(25.93±2.80)mm和(13.72±2.88)mm,完整种子平均风干重(0.0153±0.0033)g,脱翅种子平均重(0.0112±0.0024)g,种翅占整个种子的(26.56±4.53)%;伊桐种子播前采取不同处理方式的平均发芽天数、平均发芽指数和平均发芽率等存在一定或是比较明显的差异,浸种、脱翅或两者相结合的方式均能有效缩短种子发芽时间并明显提高种子发芽率;伊桐幼苗的株高和基径的月均生长量分别介于3.3～16.3cm和0.32～2.80mm之间,其中经浸种处理的前期生长较快,未经浸种处理的中期生长较快。     

Effect of Geographic Origin and Ex Situ Growing Site on Phenology,Morphology, and Seed Yield of Yarrow (Achillea millefolium L.) Germplasm from the Rhaetian Alps,Italy

Species adapted to prevailing soil and climatic conditions and native to the same geographic context are increasingly recommended for ecological restoration at high altitude. Better knowledge is required on the level of variation within these species for morpho‐physiological traits and seed yield. Adequate and affordable seed production is a prerequisite for native species to be widely adopted for restoration interventions. This study evaluated the variation of yarrow (Achillea millefolium L.) germplasm collected in the Rhaetian Alps, Italy. The worth of yarrow for restoration at high altitude has been repeatedly noted. The main goal of the study was to identify promising materials for selection purposes. The explored mountain section harbored valuable variation for traits of interest. Altitude, climate, and soil should frame an ecological correspondence between the collection site and the target restoration site, to secure the value of the native germplasm outside the boundaries of its collection area. The work also assessed any interaction between the germplasm and the ex situ growing environments, represented by a mountain and a lowland site, and verified the feasibility of the latter for seed multiplication. The growing site remarkably affected several morpho‐physiological characters. The lack of difference in mean seed yield between the two altitude‐contrasting sites suggested that seed multiplication of yarrow could also be conveniently carried out in lowland environments, with advantages in terms of production costs. Nonetheless, different yield responses in the two sites were observed, emphasizing the need of choosing the material to multiply based on the environment adopted for seed production.     

Seed‐enhancement combinations improve germination and handling in two dominant native grass species

The use of native grasses for both restoration and commercial purposes is becoming increasingly important globally. Many native grasses have limited success in seed‐based restoration (e.g. post‐mine rehabilitation) and commercial industries (e.g. agriculture) due to poor seed germination and handling. Seed‐enhancement technologies can assist in overcoming these barriers. This study aimed to use combinations of seed enhancements to overcome the germination and handling challenges in two dominant Australian native grass species with demand in restoration and commercial industries (Triodia wiseana C.A Gardner and Rytidosperma caespitosum [Gaudich.] Connor & Edgar). Selected enhancements included hydropriming (including inoculation with karrikinolide [KAR₁]), flash flaming, and seed coating. Combinations of these seed enhancements allowed improvements in both germination (by up to 55% and 18% for T. wiseana and R. caespitosum, respectively) and floret geometry (flowability, as measured through a mechanized seeder, improved by up to 6‐fold and 17‐fold for T. wiseana and R. caespitosum, respectively), with the order of enhancement application being important. The responses of each species to enhancements corresponded with key biological processes and ecological cues required for recruitment events in nature, such as fire and rainfall events. Triodia wiseana germination was driven by fire‐related cues (i.e. KAR₁, flaming), while R. caespitosum germination was highest in response to moisture‐related cues (i.e. hydro‐priming). Responses to seed enhancements (and combinations of) may have implications for the management and scaled use of the targeted species. This can assist in improving the restoration and commercial success of the study species, and potentially other grasses with germination and handling challenges, into the future.     

The response of ecosystem CO<Subscript>2</Subscript> exchange to small precipitation pulses over a temperate steppe

We evaluated the process of salt-marsh colonization in early successional stages of salt-marsh restoration and investigated how the sequence of species establishment related to different success factors. Vegetation data were collected by permanent plots from the restoration site and adjacent, reference salt marshes during three consecutive periods. Seed length, width and mass were used as dispersal traits, and Ellenberg moisture, salinity and nutrient indices as indicators of site suitability. Seed production in the reference site and seed bank in the restoration site were also investigated. The establishment of salt-marsh species within the restoration site was rapid (less than 5 years). The cover of plant species was not correlated between the restored and the reference sites at the first year of restoration, but this correlation was significant during the following years. Seed availability was more important in explaining the sequence of species establishment than salt and nutrient-limitation tolerance. The first colonizers are known as massive seed producers, with shorter seed length and lower seed mass, which probably increased buoyancy. Among dispersal and site traits, seed length and mass, and in a less extent salinity and nutrients, indicated a relationship with new colonizers. Despite few species have not (yet) appeared in vegetation and seed bank in the restoration site, the existence of an existing salt marsh adjacent to the restoration site is shown to be vital for fast colonization of newly created intertidal areas.     

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.     

Polyploidy: a missing link in the conversation about seed transfer of a commonly seeded native grass in western North America

The use of local, native plant materials is now common in restoration but testing for polyploidy in seed sources is not. Diversity in cytotypes across a landscape can pose special seed transfer challenges, because the methods used to determine genetically appropriate materials for seed transfer do not account for cytotypic variation. This lack of consideration may result in mixing cytotypes through revegetation, which could reduce long‐term population viability. We surveyed nine populations of a native bunchgrass, Pseudoroegneria spicata, in three EPA Level III Ecoregions in the western United States to determine the frequency of polyploidy, whether there are differences in traits (phenotype, fecundity, and mortality) among plants of different cytotypes, and whether cytotype frequency varies among ecoregions. We assessed trait variation over 2 years in a common garden and determined ploidy using flow cytometry. Polyploidy and mixed cytotype populations were common, and polyploids occurred in all ecoregions. Four of the nine populations were diploid. The other five had tetraploids present: three had only tetraploid individuals whereas two had mixed diploid/tetraploid cytotypes. There was significant variation in traits among cytotypes: plants from tetraploid populations were larger than diploid or mixed populations. The frequency and distribution of cytotypes make it likely that seed transfer in the study area will inadvertently mix diploid and polyploid cytotypes in this species. The increasing availability of flow cytometry may allow ploidy to be incorporated into native plant materials sourcing and seed transfer.     

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.     

The interacting effects of herbivore exclosures and seed addition in a wet meadow

Seed limitation may prevent successful restoration of native plant communities. Seed addition is a common restoration practice but the role of small mammals in affecting seedling recruitment is not well understood. The purpose of this investigation was to test the relative effect of seed introduction in combination with small mammal and bird exclosures in an Ohio wet meadow. We ask whether the ambient population of Microtus pennsylvanicus (1) alters species composition (e.g. forb/grass/sedge, invasive, non-native); (2) influences plant diversity; and (3) reduces the effect of increasing local plant richness through seed introductions. We established a 2×2 factorial design including a seed addition treatment (0 and 20 seed species added) and an exclosure treatment (open and fenced to exclude all mammalian and bird herbivores and granivores). Seeds from twenty native species were selected to represent a broad range of plant life forms typically found in temperate eastern North American wet meadow communities. All species were obligate or facultative wetland species with forbs, grasses and sedges represented. We found that forb species increased inside exclosures, especially in the seed addition treatment. We also found that relative biomass of invasive species was reduced in exclosures and with seed addition. Species richness increased with seed addition; however, exclosures significantly increased species richness and diversity, particularly of those species that were experimentally introduced by seed. Our results support the seed limitation hypotheses. It is also evident that seed and seedling predation are important factors that can control wet meadow community composition and diversity.     

Should I pick that? A scoring tool to prioritize and valuate native wild seed for restoration

Commercial sources of native seed are often unavailable for ecological restoration projects or do not have a suitable provenance. Local collection of wild seed is an option, but it can be challenging to collect seed for a variety of species and set fair seed prices. Our aim was to quantify the relative effort to collect, clean, store, and propagate seed to better prioritize species and assess the value of their seed. For 57 species native to the Canadian subarctic and typical of upland habitats, we evaluated 13 poorly correlated attributes in the field and lab or using the literature. For collection attributes, regional occurrence, local abundance, seed collection rate, and collection window were normally or log‐normally distributed. Most species were easy to identify and posed few collection obstacles. Cleaning effort was evenly distributed across species and the majority could be cleaned to more than 95% purity. We only encountered orthodox seed and most species had seed longevity exceeding a year. Seed viability mostly exceeded 80%, pre‐treatment requirements were evenly distributed and the majority of species could be germinated under standard conditions. We propose a standard worksheet, in which we assign relative effort scores to the distribution of each attribute. We illustrate this approach for the revegetation planning of a remote mine site. We also propose a seed lot certificate to ensure high seed quality. This tool can be applied to various restoration applications to assess relative effort, to plan and prioritize species for restoration projects and to help set fair seed pricing.     

Living in an urban pod: Seed predation and parasitism of bruchid beetles in a native tree species

1. Urbanisation is one of the main drivers of insect species loss worldwide. However, its impacts on ecological interactions involving insects still deserve further research, especially seed predation and parasitism of seed predators.
2. Here, we evaluated the seed predation rate by the specialist bruchid beetle Pseudopachymerina spinipes and its parasitism rate in the native tree Vachellia caven (Fabaceae) along an urbanisation gradient in Cordoba (Argentina). Since resource availability can influence these ecological interactions, we also investigated whether seed and prey availability could affect seed predation and parasitism rates, respectively.
3. We sampled trees in 10 sampling sites along an urbanisation gradient estimated by the Normalised Difference Vegetation Index (NDVI) within a 100 m radius. In our system, sites with low NDVI, beyond representing the low amount of vegetation cover, also indicate high surface temperature and low availability of host trees.
4. Seed predation in V. caven and the parasitism rate of P. spinipes were significantly reduced with increasing urbanisation. Notably, seed availability at the pod level did not affect seed predation rate, while prey availability was negatively correlated with parasitism rate.
5. These findings suggest a deleterious effect of urbanisation on the studied antagonistic interactions, giving no support to the idea of resource limitation effects.