Predicting plant performance for the ecological restoration of grasslands: the role of regenerative traits

Active restoration of grassland ecosystems often requires the addition of species as seeds. However, knowledge of seed germination and initial performance of native species are still lacking in many regions, including the Brazilian grasslands. Regenerative traits (such as seed traits) have the potential to predict performance parameters related to the initial establishment of a species in a given environment. This study aimed to test the potential of regenerative traits as predictors of the performance of grassland species, information which may be useful for ecological restoration purposes. Using 15 grassland species, we carried out three parallel experiments at different conditions: cabinet, greenhouse, and field. Three performance parameters were evaluated: final germination percentage (FGP), germination rate index (GRI), and survival rate. The suitability of seed traits (mass, area, and shape) as predictors of performance parameters was evaluated for each experiment using linear regression models. Performance parameters varied among the species and experiments, but seed mass and shape slightly influenced GRI. Seed area had a positive effect on GRI, and there was a significant interaction between seed circularity and mass, which improved the GRI of round, heavy seeds. FGP and survival rate were not predicted by any of the traits. Our study highlights the importance of testing fitness–trait relationships under field conditions, which may reflect the real challenges species will face in restoration projects. Our results reinforce the need of considering regenerative traits and interaction factors to improve the understanding and prediction of the initial performance of species.     

Seed use in the field: delivering seeds for restoration success

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

生物结皮对5种不同形态的荒漠植物种子萌发的影响

生物结皮广泛分布于干旱、半干旱区, 强烈影响着土壤表层理化特性, 进而对种子散布、萌发和定居产生影响。目前关于生物结皮与植物种子萌发关系的研究结论存在争议。该文通过室内人工控制实验, 研究了生物结皮对古尔班通古特沙漠5种具不同种子形态特征的荒漠植物白梭梭(Haloxylon persicum)、蛇麻黄(Ephedra distachya)、角果藜(Ceratocarpus arenaarius)、涩芥(Malcolmia africana)和狭果鹤虱(Lappula semiglabra)的种子萌发的影响。结果表明, 在干燥和湿润两种条件下, 生物结皮对不同形态植物种子萌发均具有不同的作用。在干燥条件下, 生物结皮显著抑制了角果藜和涩芥种子的萌发(p<0.05), 对其它3种植物无显著影响; 而湿润条件下, 生物结皮显著抑制了白梭梭、角果藜和狭果鹤虱种子的萌发(p<0.05), 对蛇麻黄、涩芥则无显著影响。     

Innovative Techniques for Large‐scale Seagrass Restoration Using Zostera marina (eelgrass) Seeds

The use of Zostera marina (eelgrass) seeds for seagrass restoration is increasingly recognized as an alternative to transplanting shoots as losses of seagrass habitat generate interest in large‐scale restoration. We explored new techniques for efficient large‐scale restoration of Z. marina using seeds by addressing the factors limiting seed collection, processing, survival, and distribution. We tested an existing mechanical harvesting system for expanding the scale of seed collections, and developed and evaluated two new experimental systems. A seeding technique using buoys holding reproductive shoots at restoration sites to eliminate seed storage was tested along with new techniques for reducing seed‐processing labor. A series of experiments evaluated storage conditions that maintain viability of seeds during summer storage for fall planting. Finally, a new mechanical seed‐planting technique appropriate for large scales was developed and tested. Mechanical harvesting was an effective approach for collecting seeds, and impacts on donor beds were low. Deploying seed‐bearing shoots in buoys produced fewer seedlings and required more effort than isolating, storing, and hand‐broadcasting seeds in the fall. We show that viable seeds can be separated from grass wrack based on seed fall velocity and that seed survival during storage can be high (92–95% survival over 3 months). Mechanical seed‐planting did not enhance seedling establishment at our sites, but may be a useful tool for evaluating restoration sites. Our work demonstrates the potential for expanding the scale of seed‐based Z. marina restoration but the limiting factor remains the low rate of initial seedling establishment from broadcast seeds.     

Global patterns in seed germination after ingestion by mammals

1. Mammals play an important role in seed germination through the ingestion of fruits and seeds. Since seed germination is a basic step in seedling recruitment, understanding how mammals affect germination improves our understanding of the effect of loss of mammal populations on the dynamics of plant communities.
2. We used meta-analytical methods to describe global patterns in the effect of seed ingestion by mammals on seed germination success and rate. We collected data from 154 studies that included 115 mammal species and 448 plant species.
3. Our results showed a positive cumulative effect of mammals on seed germination. However, this effect differed between mammalian orders; thus, some groups such as elephants, primates, and new world marsupials emerged as important enhancers of seed germination. Also, the effect varied depending on the plant family and the bioregion. Increased seed germination after ingestion was positively related to fast germination.
4. This meta-analysis, the first to synthesise and compare most of the information presently available on how mammals affect seed germination after ingestion, shows a global positive effect of mammals as enhancers of seed germination. However, behind that positive effect lies a diversity of neutral, negative, and positive effects of different magnitudes, which may have multifactorial explanations. We hope that the patterns presented here open up new questions and help guide future research efforts.

     

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.     

青藏高原东缘61种常见木本植物种子萌发特性及其与生活史的关联

为了评价亚高山地带木本植物种子萌发对策及其与生活史关联, 该文对青藏高原东缘61种常见木本植物的种子萌发特征以及种子萌发与种子大小、扩散方式、海拔及生境的关系进行了调查分析。结果表明: 1) 61种植物的种子萌发率呈现偏斜、双峰分布。6种植物(9.8%)的种子萌发率大于80%, 13种(21.3%)的萌发率在60%~80%之间, 9种(14.8%)的萌发率在40%~60%之间, 8种(13.1%)的萌发率在20%~40%之间, 25种(41%)的萌发率小于20%, 其中8种植物的种子萌发率为零, 显示了该生境中木本植物的种子以休眠、低萌发率或连续稳定的萌发对策占优势, 以避免不利的环境因素。2)种子扩散方式是决定61种木本植物种间萌发率变异(R² = 24.1%, p < 0.001) 和萌发开始时间变异(R ²= 21.3%, p < 0.01) 的主要因素; 种子大小、母体植株的海拔和生境对种子萌发率和萌发开始时间几乎没有影响(p > 0.05); 风扩散的种子比脊椎动物扩散和无助扩散的种子有更高的萌发率(F = 9.219, p < 0.001) 和较早的萌发开始时间(F = 6.772, p < 0.01)。说明植物生活史特征如种子萌发与扩散方式之间存在固有而紧密的联系, 扩散能力强的种子(如风扩散)由于能散布到较远而空旷的生境, 可能避免了各种有害因素或个体竞争, 因而种子以较强的萌发能力进行拓殖; 扩散能力弱的种子(如无助扩散)由于其散布距离较近, 种子以休眠或降低萌发的方式来避免个体或同胞竞争以及各种有害因素, 以获得最大的生态利益并确保物种延续。     

六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

Seed depredation negates the benefits of midstory hardwood removal on longleaf pine seedling establishment

Midstory hardwoods are traditionally removed to restore longleaf pine on fire‐excluded savannas. However, recent evidence demonstrating midstory hardwood facilitation on longleaf pine seedling survival has brought this practice into question on xeric sites. Also, midstory hardwoods could facilitate longleaf pine seedling establishment, as hardwood litter may conceal seeds from seed predators or improve micro‐environmental conditions for seedling establishment. However, little is known about these potential mechanisms. In this study, we tracked longleaf pine seed depredation and germination in artificially seeded plots (11 seeds/m²) in a factorial design fully crossing hardwood retention or removal with vertebrate seed predator access or exclusion in the Sandhills Ecoregion of North Carolina, U.S.A. Seed depredation averaged 78% across treatments and was greatest in unexcluded plots. Hardwood retention did not affect seed depredation. Longleaf pine averaged 3.6 germinants/4 m² across treatments, and was six times more abundant where vertebrates had been excluded. Hardwood removal had a strong positive effect on seedling germination, likely due to the removal of litter, but only when vertebrates were excluded. Our results indicated midstory hardwoods are not facilitating longleaf pine seedling establishment. Nevertheless, our results indicated that hardwood removal may not increase longleaf pine seedling establishment, as seed depredation diminished the effectiveness of hardwood removal under mast seed availability. Collectively, these results demonstrate the underlying complexity of the longleaf pine ecosystem, and suggest that planting may need to be part of the restoration strategy on sites where seed depredation limits longleaf pine natural regeneration.     

Effects of temperature and storage length on seed germination and the effects of light conditions on seedling establishment with respect to seed size in Ligularia virgaurea

Ligularia virgaurea is widely distributed in the alpine meadows of the eastern Qinghai-Tibet plateau. We studied the effects of temperature and seed storage length on seed germination in a laboratory experiment, the effects of seed mass and light intensity on seed emergence in a pot experiment, and the effect of meadow disturbance intensity on seed emergence in a field experiment. Our results showed that seeds of L. virgaurea germinated well under a wide range of temperatures. Germination percentage decreased with increased seed storage length. When seeds were stored for either 6 or 12 months the germination percentage increased with decreasing temperature. The emergence percentage of large seeds was higher than that of small seeds. Seedling survival of large seeds was greater than that of small seeds in 75% and 50% of natural irradiance. Large seeds of L. virgaurea were more successful colonizers in the alpine meadows. Seeds of L. virgaurea germinated in both disturbed meadows and non-disturbed meadows, but seed emergence and seedling establishment were higher in the disturbed meadows, that is, disturbance could facilitate the recruitment and establishment of L. virgaurea .     

Cracking the case: Seed traits and phylogeny predict time to germination in prairie restoration species

Traits are important for understanding how plant communities assemble and function, providing a common currency for studying ecological processes across species, locations, and habitat types. However, the majority of studies relating species traits to community assembly rely upon vegetative traits of mature plants. Seed traits, which are understudied relative to whole‐plant traits, are key to understanding assembly of plant communities. This is particularly true for restored communities, which are typically started de novo from seed, making seed germination a critical first step in community assembly and an early filter for plant establishment. We experimentally tested the effects of seed traits (mass, shape, and embryo to seed size ratio) and phylogeny on germination response in 32 species commonly used in prairie grassland restoration in the Midwestern USA, analyzing data using time‐to‐event (survival) analysis. As germination is also influenced by seed dormancy, and dormancy break treatments are commonly employed in restoration, we also tested the effects of two pretreatments (cold stratification and gibberellic acid application) on time to germination. Seed traits, phylogeny, and seed pretreatments all affected time to germination. Of all traits tested, variables related to seed shape (height and shape variance) best predicted germination response, with high‐variance (i.e., pointier and narrower) seeds germinating faster. Phylogenetic position (the location of species on the phylogenetic tree relative to other tested species) was also an important predictor of germination response, that is, closely related species showed similar patterns in time to germination. This was true despite the fact that all measured seed traits showed phylogenetic signal, therefore phylogeny provided residual information that was not already captured by measured seed traits. Seed traits, phylogenetic position, and germination pretreatments were important predictors of germination response for a suite of species commonly used in grassland restoration. Shape traits were especially important, while mass, often the only seed trait used in studies of community assembly, was not a strong predictor of germination timing. These findings illustrate the ecological importance of seed traits that are rarely incorporated into functional studies of plant communities. This information can also be used to advance restoration practice by guiding restoration planning and seed mix design.     

Protecting direct seeded grasses from herbicide application: can new extruded pellet formulations be used in restoring natural plant communities?

Restoration of native plant communities through direct seeding often experience low seedling establishment success rates, partly due to competition with invasive weed species. To improve seeding success, herbicides can be applied to control weed competition, however, this can have negative impacts on the seeded species. Activated carbon (AC) can be incorporated into newly developed seed enhancement technologies to adsorb herbicides and increase seedling tolerance. This study expands upon research completed to date, by developing new formulations of extruded pellets containing AC, aiming to provide increased protection to seeded species and increase herbicide selectivity. We tested six extruded pellet formulations, which included two pellet formula variations, and three quantities of AC, to examine the impact on emergence (without herbicide) and mortality (with herbicide) of Lolium rigidum Gaudin (annual ryegrass). Extruded pellet formulations containing a superabsorbent polymer (3%) and AC (10%) did not impede emergence (79%), in the absence of herbicide, similar to the non‐pelleted seeds (81%). This extruded pellet formulation increased seedling tolerance to Simazine (a pre‐emergent, soil applied herbicide) application, with mortality reduced from 96% in non‐pelleted seeds, and 77% in pellets containing no AC, to 22% in pellets containing AC. The results from this study demonstrate that AC extruded pelleting can be used as a restoration seeding technology by protecting seeds from the negative effects of pre‐emergent herbicide applications. Field evaluations with native seeds will mark an important step forward to ensure seed enhancement technology options, such as AC extruded pelleting, are available for restoring natural plant communities in restoration programs.     

生物土壤结皮对种子萌发和幼苗建植的影响研究进展

生物土壤结皮（BSC）在陆地生态系统中具有重要的生态地位,尤其是旱地生态系统中,BSC占据了种子植物之间的广阔地面。因此,BSC的发展必然影响种子植物更新过程与植被空间格局;但其作用方式、影响程度等因相关研究涉及多气候要素、土壤类型、BSC组成物种和种子植物物种的差异及其不同组合,导致目前的研究结论存在广泛争议。研究综合论述了BSC改变地表微形态对种子传播过程的影响;BSC改变土壤特性（物理、化学、生物学）对种子萌发和幼苗存活与建植等关键环节的影响;并结合种子形态特征及种子萌发、幼苗建植的性状等,综合分析了BSC对种子传播、种子萌发与幼苗建植等更新过程的潜在影响机理;探讨了目前研究矛盾性结论产生的原因。总体来说,深入研究并全面揭示BSC对种子植物更新过程的影响,应加强学科交叉,将分子生物学、植物生理学、生物化学等微观研究,与遥感、野外生态因子过程监测、控制实验等宏观、中观研究结合,从机理到过程方面动态研究BSC对种子植物更新过程的影响,并引入水文模型、气候模型、种群动态模型等模型预测方法,研究气候变化、各类干扰频发的情景下,BSC对种子传播、萌发及幼苗建植过程的潜在影响,以期促进对BSC与种子植物间相互关系的研究,加深对干旱脆弱生态系统植被发展规律的认识。     

Desiccation,Moisture Content and Germination of Zostera marina L. Seed

Zostera marina is the only seagrass species whose seeds have been successfully used in large‐scale restoration. Although progress has been made in refining Z. marina restoration protocols, additional information on Z. marina seed physiology is necessary as the science of seagrass restoration evolves. We tested the germination rates of Z. marina seeds under different relative humidities and temperatures for different periods of time. Z. marina seed moisture content (MC) and germination rates were also tested when seeds were exposed to a temperature of 25°C and relative humidity of 50%. Z. marina seeds suffered higher mortality when exposed to lower relative humidity and higher temperature for longer period of exposure time. A significant negative correlation was detected between seed germination rate and MC. Z. marina seeds are sensitive to desiccation exposure and long periods of exposure to air should be prevented to minimize seed mortality when seeds are used in restoration projects.     

模拟水淹对三峡库区常见一年生草本植物种子萌发的影响

根据三峡水库水位运行时间, 设计了30、75、115、155、195和240天共6个水淹时间梯度(T-1、T-2、T-3、T-4、T-5和T-6), 采用模拟水淹的方法, 研究了不同水淹时间对三峡消落带4种常见的一年生草本植物稗(Echinochloa crusgali)、金狗尾草(Setaria pumila)、马唐(Digitaria sanguinalis)和荩草(Arthraxon hispidus)种子萌发的影响。结果表明: 1)随着水淹处理时间的增长, 这4种植物的萌发率基本上呈现先增高后降低的趋势。稗和荩草在T-1-T-5的种子萌发率显著高于对照(CK) (p < 0.05), T-6和CK之间差异不显著。金狗尾草T-2、T-3的萌发率显著高于CK (p < 0.05), T-1、T-4-T-6与CK之间无显著性差异。马唐在水淹处理(T-1-T-6)的萌发率均显著高于对照(CK), 但长时间水淹处理(T-4-T-6)对萌发的促进作用要低于短时间水淹处理(T-1-T-3)。这说明一定时间的水淹有利于打破种子休眠并提高种子萌发率。2)一定时间的水淹处理加快了稗、金狗尾草、马唐和荩草的萌发进程。对照组种子的日萌发率较均匀, 萌发曲线较平缓。而水淹处理的种子多集中在3-5天内大量萌发。和对照相比, 一定时间的水淹处理显著提高了这4种植物种子的萌发指数, 缩短了种子的萌发持续时间, 提早了种子萌发高峰时间和达到50%萌发率的时间。长时间的水淹对种子的萌发进程影响不大。3)总体来说, 稗、金狗尾草、马唐和荩草在各个处理下的萌发率均较高(> 40%), 可以考虑作为三峡消落带植被恢复的备选物种。     

Priming effects on seed germination in Tecoma stans (Bignoniaceae) and Cordia megalantha (Boraginaceae), two tropical deciduous tree species

Successful revegetation necessarily requires the establishment of a vegetation cover and one of the challenges for this is the scarce knowledge about germination and seedling establishment of wild tree species. Priming treatments (seed hydration during a specific time followed by seed dehydration) could be an alternative germination pre-treatment to improve plant establishment. Natural priming (via seed burial) promotes rapid and synchronous germination as well as the mobilisation of storage reserves; consequently, it increases seedling vigour. These metabolic and physiological responses are similar to those occurring as a result of the laboratory seed priming treatments (osmopriming and matrix priming) applied successfully to agricultural species. In order to know if natural priming had a positive effect on germination of tropical species we tested the effects of natural priming on imbibition kinetics, germination parameters (mean germination time, lag time and germination rate and percentage) and reserve mobilisation in the seeds of two tree species from a tropical deciduous forest in south-eastern México: Tecoma stans (L Juss. Ex Kunth) and Cordia megalantha (S.F Blake). The wood of both trees are useful for furniture and T. stans is a pioneer tree that promotes soil retention in disturbed areas. We also compared the effect of natural priming with that of laboratory matrix priming (both in soil). Matrix priming improved germination of both studied species. Natural priming promoted the mobilisation of proteins and increased the amount of free amino acids and of lipid degradation in T. stans but not in C. megalantha. Our results suggest that the application of priming via the burial of seeds is an easy and inexpensive technique that can improve seed germination and seedling establishment of tropical trees with potential use in reforestation and restoration practices.     

Are fungi important for breaking seed dormancy in desert species? Experimental evidence in Opuntia streptacantha (Cactaceae)

Seeds of Opuntia spp. have physiological dormancy; they need a period of after-ripening to break dormancy, and the embryos have low growth potential. We evaluated the combined effects of seed age and presence of fungi on the testa on germination of Opuntia streptacantha, an abundant species in the Chihuahuan Desert (Mexico), assuming that older seeds have broken seed dormancy and fungi can reduce mechanical resistance to germination. In a preliminary experiment, we found no germination of 9-year-old (1998) and freshly collected (2007) seeds. However, we obtained 67% and 27% germination from 9-year-old and fresh non-sterilized seeds, respectively, and found fungi growing on the testa of all germinated seeds. Two fungal strains were isolated and identified using ribosomal internal transcribed spacer (ITS) sequence analysis: Penicillium chrysogenum and Phoma sp. In a second experiment, we inoculated seeds with strains of P. chrysogenum and Phoma sp., as well as Trichoderma koningii and binucleate Rhizoctonia (Gto17S2), to evaluate their ability to break seed dormancy. Seeds inoculated with P. chrysogenum, Phoma sp. and T. koningii had higher germination than controls for both seed ages, but germination was higher in older seeds. Scanning electron microscopy showed that these fungi eroded the funiculus, reducing its resistance. Binucleate Rhizoctonia did not lead to germination and controls had almost no germination. Our results strongly indicate that fungi are involved in breaking seed dormancy of O. streptacantha, and that the effect of fungi on seeds is species-specific.     

种皮和环境温度对橡胶树种子萌发的影响

在人工气候箱控制的条件下,研究了种皮和环境温度对橡胶树(Hevea brasiliensis)种子萌发的影响.结果表明,种皮对橡胶树种子的吸水和萌发具有明显的机械限制作用;去除种皮后种子萌发速率加快,萌发指数增大,但萌发率从83.8%下降到47.8%,幼苗活力指数从332.3下降到187.9.15℃以下的温度明显推迟种子萌发,且萌发率下降.去除种皮后的种仁萌发率、萌发指数和幼苗活力指数均随着温度的升高而增大;有种皮种子的萌发率、萌发指数和幼苗活力指数分别在20℃(94.7%)、30℃(1.86)和25℃(374.4)时最大,而萌发速率系数随着温度的升高先增大后减小.在橡胶树栽培和胶园建设中,可利用种皮对种仁水分散失的限制作用以及对低温的不敏感性,有效延长橡胶树种子的贮藏时间,提高萌发率和籽苗质量.     

Transfer seeds,hay, or soil blocks? The importance of the completeness of biological inputs to address dispersal and establishment limitations during the restoration of plant assemblages in floodplain grasslands

This study investigated different techniques of grassland restoration to overcome dispersal or establishment limitation, which are key processes influencing early-successional plant community assembly. A fully randomized in situ experiment was set up in a former arable land in a floodplain along the Garonne river (south-western France) to test for the effect of (1) the type and completeness of the biological input (any biological material—seed, hay, and soil—transferred to the site under restoration), (2) soil disturbance by deep tillage, and (3) their interaction on plant community dynamics for 5 years. All inputs influenced the plant community structure and composition. The effects depended on the type of the input but not necessarily in link with its completeness. High density and diversity seed mixture led to high levels of richness and relative abundance of target species like soil blocks, the input considered as the most complete. During the first year, hay transfer mostly influenced community assembly through negative litter effects. Delayed germination of several species contributed to buffer these early effects. This study supports the importance of dispersal limitation during early succession in degraded grassland ecosystems whereas soil disturbance had only subtle effects on the seed bank and standing plant community, indicating that competition and establishment limitation were of secondary importance in our study system. Our results suggest that even low-frequency immigration events (e.g. when a few seeds are transferred within hay) can be of great importance providing that a sufficiently long time period is considered for grassland restoration.     

Effect of seed passage through vertebrate frugivores'' guts on germination: a review

The capacity of seeds to germinate after ingestion by frugivores is important for the population dynamics of some plant species and significant for the evolution of plant-frugivore interactions. In this paper the effects of different vertebrates on seed germination of nearly 200 plant species are reviewed, searching for patterns that predict the circumstances in which germination of seeds is enhanced, inhibited, or unaffected by the passage through the digestive tract of a seed disperser. It was found that seed dispersers commonly have an effect on the germinability of seeds, or on the rate of germination, or both, in about 50% of the plants they consume, although the diversity of animal species tested so far is still rather low (42 bird species, 28 non-flying mammals, 10–15 bats, 12 reptiles, 2 fishes). Enhancement of germination occurred about twice as often as inhibition.

In spite of the morphological and physiological differences in their digestive tracts, the different animal groups tested have similar effects on seed germination, although non-flying mammals tend to influence germination slightly more often than the other groups. Data on fishes are still too scarce for any generalization. Seed retention time in the dispersers' digestive tract is one factor affecting germination, and helps to explain the variation in seed responses observed among plant species, and even within a species. However other factors are also important; for example, the type of food ingested along with the fruits may affect germination through its influence on chemical or mechanical abrasion of the seed coat. Seed traits such as coat structure or thickness may themselves be responsible for some of the variation in seed retention times. Seeds of different sizes, which usually have different transit times through frugivores, and seeds of either fleshy or dry fruits, show often similar germination response to gut passage.

Seeds of different plants species differ strongly in their germination response after ingestion, even by the same frugivore species. Congeneric plants often show little consistency in their response. Even within a species variation is found which can be related to factors such as the environmental conditions under which germination takes place, seed morphology, seed age, and the season when the seeds are produced.

The effect of gut passage on germination differs between tropical and temperate zones. Seed germination of both shrubs and trees (data on herbaceous species are still scarce) in the temperate zone is more frequently enhanced than in the tropics. This result supports the hypothesis that enhanced germination may be more advantageous in unpredictable or less constant environments. Significant differences in frugivore-mediated germination are also found among different life forms. In both tropical and temperate zones, trees appear to be consistently more affected than shrubs or herbs. This might be due to an overall higher thickness of the seed coats, or to a higher frequency of seed-coat dormancy in tree species.

The influence of frugivory upon the population dynamics of a species has to be evaluated relative to other factors that influence germination and seedling recruitment at a particular site. Whether seed ingestion by dispersers is really advantageous to a plant (as has commonly been assumed) can only be assessed if we also determine the fate of the ingested seeds under natural conditions, and compare it to the fate of seeds that have not been ingested.