Rodent seed predation and seedling recruitment in mesic grassland

Seedling recruitment of two grasses (Arrhenatherum elatius and Festuca rubra) and two herbs (Centaurea nigra and Rumex acetosa) was measured in areas with and without rodents to which seeds of each species were sown at three seed densities (1000, 10,000 and 50,000 seeds m⁻²) in two seasons (spring and autumn 1995). Seed removal was measured for 10-day periods and the fate of seedlings was followed for 15 months after sowing. The proportion of seed removed ranged from 6 to 85% and increased with increasing seed density for each species. Rodents had no effect on seedling emergence or survival in the spring sowing. In the autumn sowing, rodents reduced seedling emergence of all four species sown at 1000 and 10,000 seeds m⁻² but had no impact at 50,000 seeds m⁻², presumably because of microsite limitation. We suggest the difference between spring and autumn arose because emergence was seed limited in autumn but microsite limited in spring; microsite availability was higher in autumn because a summer drought killed plants, reduced plant biomass and opened up the sward. Fifteen months after the autumn sowing, fewer A. elatius and C. nigra seedlings survived on plots exposed to rodents. This result reflected not only the reduced seedling emergence but also increased seedling mortality (seedling herbivory) in sites exposed to rodents. In contrast, F. rubra and R.acteosa showed density-dependent seedling survival which compensated for initial differences in seedling emergence, so that no effect of rodents remained after 15 months. The results suggest that rodent seed predation and seedling herbivory exert strong effects on seedling recruitment of A.elatius and C. nigra when recruitment conditions are favourable (conditions that lead to high microsite availability) and may contribute to both species being maintained at low densities in the grassland. The results also demonstrate that highly significant impacts of rodent seed predation at the seedling emergence stage can disappear by the time of plant maturation. Received: 2 March 1998 / Accepted: 28 September 1998     

Soil seed bank,seedling recruitment and actual species composition in an old and isolated chalk grassland site

A soil seed bank study was conducted in an old and isolated chalk grassland site (area ca. 0.05 ha) with a high species richness in The Netherlands, over a two-year period. Each soil seed bank sample was divided into two halves: one part was sown immediately in trays with a layer of sterilized sand and put in a greenhouse, whilst the other half received a chilling period before being sown. Plant recruitment in the field was studied during the same period in permanently marked plots. Seeds of species which had become extinct during the period 1944–1970 were not found in the soil seed bank in the late eighties. One group of species still present in the actual vegetation neither showed any signs of recruitment from seed nor had a soil seed bank. Species in this group that do not recruit vegetatively can be considered as directly threatened by extinction. A group in the actual vegetation did not shown any recruitment, but nevertheless had germinable seed present in the soil. Among this group, there are some annuals; however, the majority of the species are characterized by the potential to recruit vegetatively. The majority of the species present in the actual vegetation (45%) demonstrate seedling recruitment and are also present as seeds in the soil, although there is some quantitative discrepancy between these categories. The seed bank did not reflect former stages of the actual vegetation. Based on the results, predictions can be made on the future development of the vegetation and on the fate of a number of species. Adjustment of the management regime is aimed at maintenance of the species richness of the site. This is important for nature preservation purposes since a high number (ca. 27%) of the species in the chalk grassland of this small site are on the Dutch Red List.     

Effects of arrival order and seed size on germination of grassland plants: Are there assembly rules during recruitment?

Assembly rules provide a tool to simplify the complexity of interactions that shape ecological communities. The prime objective of this study was to examine an experimental approach to the study of assembly rules, and also to examine some premises of such rules. We investigated possible assembly rules pertaining to interactions among seeds and seedlings during germination. An experiment was made on randomly sampled species pairs from a larger pool of grassland species differing in seed size, and included treatments altering the presence and ordering of species arrival (seed sowing), and nutrient level. The basic premises suggested for the rules were that a congruence exists among species in the response of germination: (i) to environmental variation; and (ii) to the presence of other species. Both of these premises were supported. In nine of 11 investigated species, germination responded to sowing treatments. Germination was commonly enhanced in species sown after another species. Inclusion of a third species did not alter two-species interactions during germination, indicating that increasing complexity may not change fundamental rules of assembly. Seed size had no systematic effect on how species behaved in the germination treatments. We suggest that assembly rules during recruitment may be useful for understanding pattern and process in plant communities, and that a method with random sampling of two-species interactions is useful to handle the complexity of multi-species communities. However, studies under field conditions are necessary to make further advances in the formulation of realistic assembly rules.     

Effect of seed predation on seed bank size and seedling recruitment of bush lupine (Lupinus arboreus)

Whether seed consumers affect plant establishment is an important unresolved question in plant population biology. Seed consumption is ubiquitous; at issue is whether seedling recruitment is limited by safe-sites or seeds. If most seeds inhabit sites unsuitable for germination, post-dispersal seed consumption primarily removes seeds that would otherwise never contribute to the population and granivory has minimal impacts on plant abundance. Alternatively, if most seeds ultimately germinate before they lose viability, there is greater potential for seed consumption to affect plant recruitment. Of the many studies on seed consumption, few ask how seed loss affects seedling recruitment for species with long-lived seed banks. We examined post-dispersal seed predation and seedling emergence in bush lupine (Lupinus arboreus), a woody leguminous shrub of coastal grasslands and dunes in California. We followed the fate of seeds in paired experimental seed plots that were either protected or exposed to rodent granivores in grassland and dune habitats. Significantly more seeds were removed by rodents in dunes than grasslands. In dunes, where rodent granivory was greatest (65% and 86% of seeds removed from plots by rodents in two successive years), there is a sparse seed bank (6.6 seeds m⁻²), and granivory significantly reduced seedling emergence (in the same two years, 18% and 19.4% fewer seedlings emerged from exposed versus protected plots), suggesting seed rather than safe-site limited seedling recruitment. In contrast, rodents removed an average of 6% and 56% of seeds from grassland plots during the same two years, and the grassland seed bank is 43-fold that of the dunes (288 seeds m⁻²). Even high seed consumption in the second year of the study only marginally influenced recruitment because seeds that escaped predation remained dormant. Burial of seeds in both habitats significantly reduced the percentage of seeds removed by rodents. Results suggest that granivores exert strong but habitat-dependent effects on lupine seed survival and seedling emergence. Received: 24 October 1996 / Accepted: 4 February 1997     

放牧方式影响高寒草地矮生嵩草种子大小与数量的关系

种子大小和种子数量变异是植物适合度研究的核心问题, 探究不同大型草食动物对嵩草(Kobresia spp.)草地优势种种子大小与数量的影响, 有助于了解其繁殖策略和种群更新机制。该研究依托青藏高原高寒草地-家畜系统适应性管理技术平台, 对不放牧(对照)、牦牛单牧、藏羊单牧、牦牛藏羊1:2混牧、牦牛藏羊1:4混牧、牦牛藏羊1:6混牧6个放牧处理下矮生嵩草(K. humilis)的种子大小和数量特征, 种子大小和数量与生殖性状的关系, 种子大小和数量的权衡关系进行研究。结果显示: 1)放牧处理增加矮生嵩草种子大小15%以上, 增加种子数量30%以上; 除了牦牛藏羊1:2混牧处理, 其他放牧处理与对照相比种子大小变异系数降低15%以上, 种子数量变异系数降低25%以上。2) Pearson相关分析结果显示, 放牧处理下矮生嵩草种子数量、种子大小与生殖相关性状均呈正相关关系。3)放牧处理增加了矮生嵩草种子大小与数量的权衡, 单条生殖枝质量是影响种子大小与数量权衡的重要性状。研究表明, 即使是中度放牧, 家畜依旧是矮生嵩草资源获取的限制性因子; 长期的放牧改变了矮生嵩草性状间的潜在联系和权衡关系, 稳定了种子大小和种子数量特征, 并通过提高种子大小和种子数量的方式优化繁殖策略, 提高了子代的竞争力和适合度。     

Seed size,number and strategies in annual plants: a comparative functional analysis and synthesis

Background and AimsPlants depend fundamentally on establishment from seed. However, protocols in trait-based ecology currently estimate seed size but not seed number. This can be rectified. For annuals, seed number should simply be a positive function of vegetative biomass and a negative function of seed size.MethodsUsing published values of comparative seed number as the ‘gold standard’ and a large functional database, comparative seed yield and number per plant and per m² were predicted by multiple regression. Subsequently, ecological variation in each was explored for English and Spanish habitats, newly calculated C-S-R strategies and changed abundance in the British flora.Key ResultsAs predicted, comparative seed mass yield per plant was consistently a positive function of plant size and competitive ability, and largely independent of seed size. Regressions estimating comparative seed number included, additionally, seed size as a negative function. Relationships differed numerically between regions, habitats and C-S-R strategies. Moreover, some species differed in life history over their geographical range. Comparative seed yield per m² was positively correlated with FAO crop yield, and increasing British annuals produced numerous seeds. Nevertheless, predicted values must be viewed as comparative rather than absolute: they varied according to the ‘gold standard’ predictor used. Moreover, regressions estimating comparative seed yield per m² achieved low precision.ConclusionsFor the first time, estimates of comparative seed yield and number for >800 annuals and their predictor equations have been produced and the ecological importance of these regenerative traits has been illustrated. ‘Regenerative trait-based ecology’ remains in its infancy, with work needed on determinate vs. indeterminate flowering (‘bet-hedging’), C-S-R methodologies, phylogeny, comparative seed yield per m² and changing life history. Nevertheless, this has been a positive start and readers are invited to use estimates for >800 annuals, in the Supplementary data, to help advance ‘regenerative trait-based ecology’ to the next level.     

Effects of altitude and seed size on germination and seedling survival of heathland plants in north Spain

The effects of altitude and seed size on germination and seedling survival were studied in Calluna vulgaris, Erica cinerea and Erica vagans. Experiments were carried out in the laboratory over a one year period. Seeds collected from heathlands of different altitudes were divided in two size classes. They were sowed on moist filter paper inside Petri dishes which were placed in chambers at 20 °C and a photoperiod of 12 h light/12 h darkeness. The seeds of Calluna vulgaris were the first to begin germination and had the highest rate and percentage of germination. The germination of Erica vagans was moderate, while seeds of Erica cinerea germinated later and the germination was very low. Seeds of Calluna vulgaris and E. cinerea collected at the highest altitudes had the highest germination percentages. Seed size in Calluna vulgaris and E. cinerea did not affect germination. However, large seeds of Erica vagans had higher germination rates and percentages than small seeds. The large seed size of Calluna vulgaris contributes to a better survival and growth of its seedlings.     

Irrigation and fertilization effects on seed number,size, germination and seedling growth: implications for desert shrub establishment

Plants with limited resources adjust partitioning among growth, survival, and reproduction. We tested the effects of water and nutrient amendments on seed production, size, and quality in Sarcobatus vermiculatus (greasewood) to assess the magnitude and importance of changes in reproductive partitioning. In addition, we assessed interactions among the environment of seed-producing plants (adult plant scale), seed size, and seedling microenvironment (seedling scale) on successful seedling establishment. Interactions of these factors determine the scale of resource heterogeneity that affects seedling establishment in deserts. Both total number of seeds produced per plant and seed quality (weight and germination) increased significantly in the enriched treatment in a 3-year field experiment. Seedling length 3 days after germination and seed N concentration, other measures of seed quality, were higher for seed from both irrigated and enriched plants than for seed from control plants. Field S. vermiculatus seed production and quality can be substantially increased with irrigation and nutrient enrichment at the adult plant scale and this allows management of seed availability for restoration. However, based on a greenhouse study, seedling environment, not the environment of the seed-producing plant or seed size, was the most important factor affecting seedling germination, survival, and growth. Thus it appears that production of more seed may be more important than improved seed quality, because higher quality seed did not compensate for a low-resource seedling environment. For both natural establishment and restoration this suggests that heterogeneity at the scale of seedling microsites, perhaps combined with fertilization of adult shrubs (or multi-plant patches), would produce the greatest benefit for establishing seedlings in the field.     

Influences of neighbouring plants on seedling establishment in limestone grassland

Abstract. Influences of neighbouring plants on seedling establishment of six dicotyledonous species was investigated in a nutrient-poor limestone grassland (Mesobrometum) in northern Switzerland. Microsites with different vegetational structure were created and seeds sown in them. The fate of emerging seedlings was followed for one or two years. A plant surviving its first growing season was regarded as established. Shelter by neighbouring plants appeared to be necessary for the establishment of Arabis hirsuta and Primula veris. Because of drought and frost heave, hardly any seedlings of these species established in gaps (4 - 21 %). In microsites with vegetation, their survival was significantly higher (40–57 %). Neighbouring plants slightly reduced the survival of Plantago lanceolata and Sanguisorba minor, but these species established well both in gaps (74 - 81 %) and in vegetated microsites (54 - 67 %). Medicago lupulina established well in all microsites in one year (71 - 79 %), but poorly in the next year (18–32 %). Linum catharticum emerged poorly in one year and was completely extinguished by a fungal pathogen the following year. The overall conclusion is, that gaps are of minor importance for recruitment of these species in this grassland. Physical hazards and pathogens control seedling establishment to a greater extent than competition by neighbouring plants. Some species are hardly able to establish without shelter of vegetation. Seed size is an important factor for success of establishment, especially in gaps. It is suggested, that the relatively low productivity and the absence of litter accumulation (due to mowing and biomass removal) are important conditions for the observed behaviour of the seedlings and juvenile plants in this community.     

Demographic,seed and microsite limitations to seedling recruitment in semi-arid mine site restoration

Plant and Soil - Understanding limitations to plant recruitment is a key element in devising effective restoration of semi-arid ecosystems: only when these limitations are identified can management...     

A model for seed size variation among plants

Summary I developed a model for seed size variation among plants assuming that the pollen captured per flower depends on both the allocation to pollen capture mechanisms per flower and the number of flowers on each plant. I showed that the optimal seed size increases with (1) the total resource allocation to reproduction, (2) decreasing outcross pollen availability, (3) decreasing probability of seedling establishment and (4) decreasing selfing rate. However, optimal seed size does not depend on the total resource allocation if the total number of pollen grains captured by a plant increases linearly with its flower number. In addition, the optimal seed size is not always positively correlated with the optimal resource allocation to pollen capture mechanisms per flower. I discussed implications of the results for seasonal decline in seed size and seed size variations among populations, such as alutitudinal variation.     

Abundance, distribution and life histories of grassland plants: a comparative study of 81 species

1 This study examines the abundance and distribution of grassland plant species in particular relation to features affecting colonization. Seed production (inversely related to seed size) and recruitment success (positively related) affect colonization ability, suggesting that seed size can be used as a key trait.
2 Data on seed size, dispersal mode, life form, geographical range size and abundance were gathered for 81 grassland plant species in a field study area in Sweden. Seed production and plant size were estimated for 69 of these species. Analyses were performed both across species, with species treated as independent data points, and for 43 'phylogenetically independent contrasts'.
3 The cross-species analyses suggested that local abundance was related to life forms but not dispersal or plant size. Perennials were generally most abundant, as were clonal species. If abundance reflects colonization we predicted that species with intermediately sized seeds (or intermediate seed production) would be most abundant, and this was supported by the phylogenetic contrast but not by cross-species analyses. In the former analysis, a high abundance of species was significantly associated with a small seed size deviation (and seed number deviation) from the median values of these traits in the community.
4 Local abundance, seed production and seed size deviation from the community median value were positively related to geographical range size in the cross-species analysis, but no relationships were seen in the phylogenetic contrast analysis.
5 We conclude that colonization processes do have a significant influence on abundance patterns in grasslands. Seed size is a key trait for colonizing ability, and the effects of the trade-off of seed size vs. seed number must be considered. No single mechanism can be identified that influences both abundance and geographical distribution range.     

Seedling recruitment in semi-natural pastures: the effects of disturbance, seed size, phenology and seed bank

We examined recruitment patterns in semi-natural pastures and their relationships to disturbance and species composition at different spatial scales (0.01 m² up to 4 m²) of both the vegetation and the seed bank. Possible associations between seedling recruitment and phenology were also studied. The study was performed in four pastures with different management history. Seedling recruitment was generally enhanced by disturbance, with the greatest effect for small-seeded species that germinate in the autumn. The local species-pool contributed to a large extent to seedling recruitment; 83 % of the recruited species were found in the vegetation, whereas 44 % of the recruited species were found in the seed bank. There were a total of 88 species found in the vegetation at different spatial scales; 61 % were recorded at the smallest spatial scale (0.01 m²). The area sampled at this scale comprised 1.25% of the area examined, indicating a small scale structuring of diversity in the pastures. The species number in the vegetation varied between 7 and 14 for the smallest scale (0.01 m²), and between 23 and 42 for the largest scale (4 m²), in the four pastures. The species number in the seed bank was less variable. Like most studies of perennial grasslands, we found no close correspondence between species distribution in the vegetation and in the seed bank. Two of the pastures had a joint management history and exhibited a close similarity of species in the seed bank, despite that one of the pastures was abandoned already in the beginning of this century.     

The effect of mollusc grazing on seedling recruitment in artificially created grassland gaps

Two experiments conducted in spring and autumn 1992 examined the effect of mollusc grazing on seedling regeneration from natural grassland seedbanks by creating artificial gaps in plots in a grassland sward. Molluscs were excluded from half the gaps by application of molluscicide. Mollusc grazing in both the spring and autumn experiment significantly reduced seedling recruitment, though the intensity of grazing was greatest in autumn. Recruitment of five species was markedly influenced by molluscicide application. In spring, plots from which molluscs were excluded contained significantly more seedlings of Chenopodium polyspermum and Ranunculus acris. In the autumn, exclusion of molluscs resulted in increased populations of R. acris, Stellaria graminea and Rumex acetosa. Cerastium holosteoides populations were greatest in autumn grazed plots. Other species, notably the grasses Holcus lanatus and Agrostis capillaris and the legume Trifolium repens were unaffected by molluscicide application. Species diversity was significantly decreased by molluscicide application in the autumn. Gap size significantly affected the recruitment of two species. Ranunculus acris populations were significantly higher in small gaps in both spring and summer, while Chenopodium recruitment in the spring was greater in small gaps. Gap size also significantly influenced the risk of mollusc attack on Ranunculus as molluscs appeared to show an aggregative feeding response in the high seedling density small gaps. Selective grazing of vulnerable seedlings by molluscs may influence the eventual relative proportions of the species present and so provide a potent mechanism in shaping community composition in grasslands.     

The effect of management on productivity,litter accumulation and seedling recruitment in a Carpathian mountain grassland

The management regime may have a significant impact on the productivity and dynamics of grasslands, but the causal relationships influencing grassland conservation value are still not completely understood. Changes of selected community characteristics, such as standing crop, proportion of forbs in the standing crop, litter amount, litter decomposition and seedling recruitment, were investigated in a 4 year manipulative experiment in a mountain grassland in Slovakia. The aim of the research was to compare changes in newly abandoned sites and sites where restoration measures were applied after 20 years of abandonment. The sites were located in areas containing two vegetation types of the Arrhenatherion alliance (wet Poo-Trisetetum and dry Anthoxantho–Agrostietum) with different moisture regimes. The expected increase of the standing crop after abandonment was rather slow, and more pronounced towards the end of the experiment, and in the wet meadow type (~30% increase). The restoration mowing promoted forb proportions in the biomass, but it did not decrease the standing biomass in the restored grasslands. Strong litter accumulation after abandonment was observed in subsequent years after abandonment, when the amount of litter increased about 100% in abandoned plots. Decrease in litter was also significant after the start of restoration mowing (a decrease from 258 to 159 g m⁻² in wet type and from 287 to 147 g m⁻² in dry type was noted). Accumulated litter was negatively correlated to seedling recruitment (r = −0.63 at the end of the experiment). The litterbag experiment showed that the wet type has a higher rate of decomposition, with 20% more biomass decomposed during the litter-bag experiment. The experiment confirmed a negative role of litter accumulation on seedling recruitment, with the number of seedlings per m² decreasing from 413 to 321 individuals in the abandoned wet-type site. This may lead to a decrease in species richness. Mowing along with raking of mowed biomass may be a useful tool to restore degraded mountain grasslands and to remove accumulated litter from the stands.     

Negative density dependence of seed dispersal and seedling recruitment in a Neotropical palm

Negative density dependence (NDD) of recruitment is pervasive in tropical tree species. We tested the hypotheses that seed dispersal is NDD, due to intraspecific competition for dispersers, and that this contributes to NDD of recruitment. We compared dispersal in the palm Attalea butyracea across a wide range of population density on Barro Colorado Island in Panama and assessed its consequences for seed distributions. We found that frugivore visitation, seed removal and dispersal distance all declined with population density of A. butyracea, demonstrating NDD of seed dispersal due to competition for dispersers. Furthermore, as population density increased, the distances of seeds from the nearest adult decreased, conspecific seed crowding increased and seedling recruitment success decreased, all patterns expected under poorer dispersal. Unexpectedly, however, our analyses showed that NDD of dispersal did not contribute substantially to these changes in the quality of the seed distribution; patterns with population density were dominated by effects due solely to increasing adult and seed density.     

Differences in life history traits of relatedEpilobium species: Clonality, seed size and seed number

Small changes in morphology can affect the performance and functions of organisms and hence their ecological success. In modular constructed plants, contrasting growth strategies may be realized by differences in the spatial arrangement and size of shoots. Such differences change the way in which meristems and resources are assigned to various functions during the lifespan of a plant. If such changes include the capacity to spread clonally, sexual reproduction may also be affected. I compare patterns in vegetative growth and sexual reproductive traits in four allopatric species ofEpilobium which are sometimes considered as subspecies of a single polymorphic taxon. The four species differ in the location of the buds which annually renew the aerial shoot system.E. dodonaei andE. steveni do not spread clonally and are characterized by a shrub-like habit.E. fleischeri, a species occurring only in the Alps, andE. colchicum, which occurs in the upper region of the Caucasus mountains, both produce buds on horizontal roots or plagiotropic shoots. Both alpine species exhibiting clonal growth have smaller shoots, fewer fruits and smaller seeds than the lowland species. An intraspecific trade-off between seed number per fruit and seed mass is realized. Both alpine species produce more seeds per fruit at the expense of seed mass. The morphological relationship between the four species and their geographical distribution suggest that clonal growth inE. fleischeri (restricted to the Alps) andE. colchicum (restricted to the Caucasus) is adaptively associated with the stressful conditions of alpine habitats. Our results suggest that clonal growth is not necessarily correlated with reduced reproduction by seeds. The success of plants which are already established may largely depend on clonal spread, but the colonization of new habitats depends on the production of a large number of small seeds with high dispersability.