Seed size and seedling growth: differential response of Australian and British Fabaceae to nutrient limitation

Seed size is widely held to exert an important influence over plant establishment, but while large seeds are often assumed to be at an advantage in nutrient-limited conditions, there is in fact, little consistent evidence to support this hypothesis. Here, we examined the interspecific relationship between seedling growth and seed size for Australian and British Fabaceae species in nutrient solutions deficient in nitrogen, phosphorus, potassium or all nutrients combined (distilled water). The British species showed no consistent link between mean seed mass and seedling growth in nutrient-limited conditions. By contrast, all four nutrient-deficient treatments yielded a significant relationship for the Australian species. Linear regression showed that growth under balanced nutrient conditions was positively associated with growth without nutrients, although in fewer cases for the British species. We suggest that habitat-specific differences in regeneration conditions and/or evolutionary history may influence the role that seed size plays in dictating how seedlings of different species respond to nutrient shortage. We recommend caution in attempts to link traits like seed size to wider patterns of plant community ecology.     

Importance of animal and plant traits for fruit removal and seedling recruitment in a tropical forest

The traits of animals and plants influence their interaction networks, but the significance of species' traits for the resulting ecosystem functions is poorly understood. A crucial ecosystem function in the tropics is seed dispersal by animals. While the importance of species' traits for structuring plant–frugivore networks is supported by a number of studies, no study has so far identified the functional traits determining the subsequent processes of fruit removal and seedling recruitment. Here, we conducted a comprehensive field study on fruit removal by frugivorous birds and seedling recruitment along an elevational gradient in the Colombian Andes. We measured morphological traits of birds (body mass, bill width, Kipp's index) and plants (plant height, crop mass, fruit width and seed mass) which we expected to be related to fruit removal and seedling recruitment. We tested 1) which bird and plant traits influence fruit removal, and 2) whether network metrics at plant species level, functional identities of frugivores (community‐based mean trait values) and/or plant traits were the main determinants of seedling recruitment. We found that large‐bodied bird species contributed more to fruit removal than small‐bodied bird species and that small‐sized fruits were more frequently removed than large‐sized fruits. Small plant species and plants with heavy seeds recruited more seedlings than did large plants and plants with light seeds. Network metrics and functional identities of seed dispersers were unrelated to seedling recruitment. Our findings have two important implications. First, large birds are functionally more important than small birds in tropical seed‐removal networks. Second, the detected tradeoff between fruit size and seed mass in subsequent recruitment processes suggests that the adaptability of forest plant communities to a loss of large frugivores is limited by life‐history constraints. Hence, the protection of large‐bodied frugivores is of primary importance for the maintenance of diverse tropical plant communities.     

Positive relationship between fruit removal by animals and seedling recruitment in a tropical forest

Fleshy-fruited plants rely on animal frugivores to disperse their seeds, and seed removal by frugivores may leave an imprint on seedling recruitment. However, to what extent plant–frugivore interactions are related to seedling recruitment has rarely been quantified at the community level, especially in species-rich tropical forests. In this study, we tested the effect of different plant traits on fruit removal by frugivores and tested the relative importance of fruit removal, plant traits and abiotic factors for seedling recruitment. We quantified plant–frugivore interactions of 22 fleshy-fruited plant species consumed by 56 diurnal frugivore species, and counted the number of seedlings that emerged along an elevational gradient in the Colombian Andes. We measured a set of plant traits (i.e., crop size; fruit size; seed load and mass; fruit nutritional contents), estimated the density of adult plants and recorded relevant abiotic factors (light, temperature and humidity). We found that fruit removal by frugivores was positively associated with crop size, but negatively associated with fruit length and unrelated to seed load and fruit nutritional content. Seedling densities were positively related to the density of adult plants, seed mass and fruit removal by animals. We found no relationship between abiotic factors and seedling recruitment. Our results indicate that fruit abundance and morphology are important determinants of fruit removal and that fruit removal is positively associated with seedling recruitment accounting for effects of species abundance and plant traits. We conclude that plant traits shape fruit removal and seedling recruitment at the community level, while these two crucial processes of forest regeneration are directly linked by seed dispersal of animals.     

Cotyledon damage at the seedling stage affects growth and flowering potential in mature plants

Seedling herbivory is an important selective filter influencing patterns of plant community composition. Nevertheless, while many of the mechanisms governing seedling selection by herbivores are well established, the effects of tissue loss at the seedling stage on subsequent plant development are poorly understood. Here we examined how the removal of 50 or approximately 100% of cotyledon area from 7-d-old chalk grassland seedlings affected subsequent plant growth and flowering over a 100-d period. Cotyledon damage had a significant effect on growth during the establishment phase for six of the nine species. For two species, significant effects on plant growth were manifest in 100-d-old plants. Of the five species that flowered, three developed fewer inflorescences or flowered later as a consequence of cotyledon damage suffered as a seedling. Our results show that, in addition to the direct effect of herbivory on seedling mortality, more subtle sublethal effects may also influence plant establishment. Reduced growth as a result of cotyledon damage may have implications for plant competition during the establishment phase, and on subsequent reproductive success at maturity.     

Species with greater seed mass are more tolerant of conspecific neighbours: a key driver of early survival and future abundances in a tropical forest

Multiple niche‐based processes including conspecific negative density dependence (CNDD) determine plant regeneration and community structure. We ask how interspecific and intraspecific density‐dependent interactions relate to plant life histories and associated functional traits. Using hierarchical models, we analysed how such interactions affected first‐year survival of seedling recruits of 175 species in a tropical forest, and how species abundances and functional traits are related to interspecific variation in density‐dependent effects. Conspecific seedling neighbour effects prevailed over the effects of larger conspecific and all heterospecific neighbours. Tolerance of seedling CNDD enhanced recruit survival and subsequent abundance, all of which were greater among larger seeded, slow‐growing and well‐defended species. Niche differentiation along the growth–survival trade‐off and tolerance of seedling CNDD strongly correlated with regeneration success, with manifest consequences for community structure. The ability of larger seeded species to better tolerate CNDD suggests a novel mechanism for CNDD to contribute to seed‐size variation and promote species coexistence through a tolerance–fecundity trade‐off.     

植物天然更新过程中种子和幼苗死亡的影响因素

植物天然更新包括有种子搬运、种子库动态、种子萌发和幼苗定居等过程。从种子生产到幼苗定居的更新是植物生活史中最为敏感的阶段之一 ,多种因素的影响种子和幼苗的命运。其中包括 :( 1 )动物取食或病原体侵袭。种子在扩散和搬运过程中 ,易被小哺乳动物或无脊椎动物取食。蛀虫也可以使种子失去萌芽能力。病原体感染种子和幼苗 ,容易引起种子和幼苗的死亡。 ( 2 )异质生境的影响。在不同生境中 ,光照条件、土壤水分和化学成分等因子的组合严重影响种子和幼苗的命运。 ( 3 )干扰的影响。小尺度和大尺度的干扰都可以影响到植物更新时种子和幼苗的命运。林窗作为特殊的干扰体系 ,为不同种类植物提供了更新的机会。 ( 4 )繁殖体特征。种子大小、质量和保护色等特征影响种子和幼苗在更新过程中的生存。种子休眠期间 ,由于生理衰老和腐烂的原因使种子失去活力而不能萌发。 ( 5 )密度和距离制约。母株附近由于密度竞争的影响 ,种子和幼苗死亡率都较高。     

Seed size predicts global effects of small mammal seed predation on plant recruitment

Recent studies demonstrate that by focusing on traits linked to fundamental plant life‐history trade‐offs, ecologists can begin to predict plant community structure at global scales. Yet, consumers can strongly affect plant communities, and means for linking consumer effects to key plant traits and community assembly processes are lacking. We conducted a global literature review and meta‐analysis to evaluate whether seed size, a trait representing fundamental life‐history trade‐offs in plant offspring investment, could predict post‐dispersal seed predator effects on seed removal and plant recruitment. Seed size predicted small mammal seed removal rates and their impacts on plant recruitment consistent with optimal foraging theory, with intermediate seed sizes most strongly impacted globally – for both native and exotic plants. However, differences in seed size distributions among ecosystems conditioned seed predation patterns, with relatively large‐seeded species most strongly affected in grasslands (smallest seeds), and relatively small‐seeded species most strongly affected in tropical forests (largest seeds). Such size‐dependent seed predation has profound implications for coexistence among plants because it may enhance or weaken opposing life‐history trade‐offs in an ecosystem‐specific manner. Our results suggest that seed size may serve as a key life‐history trait that can integrate consumer effects to improve understandings of plant coexistence.     

Impacts of seedling herbivory on plant competition and implications for species coexistence

Background and Aims

Although the causes and consequences of seedling herbivory for plant community composition are well understood, the mechanisms by which herbivores influence plant species recruitment to the established phase remain less clear. The aim was to examine how variation in the intensity of seedling herbivory interacts with growth-defence trade-offs and herbivore feeding preferences to affect plant community development.

Methods

Using 14-d-old seedlings of Trifolium pratense and T. repens, relative growth and susceptibility to herbivory by the snail Helix aspersa was quantified to elucidate putative growth-defence trade-offs for these species. Then mixed assemblages of 14-d-old Trifolium seedlings were exposed to herbivory by zero, two, five or ten snails and determined how variation in the intensity of herbivory affected competitive interactions into the mature phase (as measured by total plant biomass at 120 d old).

Key Results

In the absence of herbivory, communities were dominated by T. pratense; a result expected on the basis that it yielded larger and presumably more competitive seedlings. However, when seedlings were exposed to herbivory, the balance of competition shifted. At low levels of herbivory (two snails), both Trifolium species contributed equally to total plant biomass. More intense herbivory (five snails) resulted in almost total mortality of T. pratense and dominance of the mature community by T. repens. The most intense herbivory (ten snails) effectively removed all seedlings from the experimental community.

Conclusions

The study illustrates a mechanism whereby spatio-temporal fluctuations in seedling herbivory, when coupled with species-specific variation in competitive ability and sensitivity to herbivore attack, can differentially influence plant recruitment into the mature phase. This mechanism may be a key element in our attempts to understand plant species coexistence, since fluctuations in plant recruitment are fundamental to the many theories that view coexistence as a consequence of a spatio-temporal lottery for dominance over regeneration micro-sites.Key words: Growth-defence trade-off, lottery models, plant–animal interactions, plant size variability, seedling acceptability, seedling defence, spatio-temporal niches, Trifolium pratense, Trifolium repens     

Influence of seed size on performance of non-native annual plant species in a novel community at two planting densities

Climate warming enables plant species to migrate to higher latitudes and altitudes. Within Europe, the Mediterranean harbours many species that might expand their ranges towards Western Europe. Small seed size may facilitate dispersal, however, it may impair establishment of the range-expanding plant species in the novel vegetation. In a greenhouse experiment, we examined effects of average seed size of Mediterranean plant species on their establishment in a mixed community of Western European plant species. Applying two levels of densities of the natives and a herbivory treatment, we tested how seed size is linked to response in plant growth and fitness in novel vegetation. While all non-native plant species showed a negative response to increased planting density, species with small seeds showed a less negative response. This effect persisted under herbivory. Our data suggest that small-seeded non-native plant species may tolerate competitive pressure from novel plant communities better than large-seeded species, so that small seed size may confer a higher probability of establishment of non-native species in novel communities.     

植物种子大小与幼苗生长策略研究进展

种子大小和幼苗更新对策是植物生活史策略的重要组成部分.本文从不同侧面综述了当前国内外关于种子大小和幼苗之间关系的生态学研究,包括:种子大小对萌发出土的影响,种子大小与幼苗形态生长特性之间的关系,种子大小与幼苗存活、竞争能力之间关系等,并对今后的相关研究进行了展望.在不同的微环境和植被类型下,种子大小与幼苗生长之间的关系可能有所差异;种子大小对植物幼苗生长的影响导致种子大小不同的植物对植被幼苗更新补充的贡献不同;种子大小与幼苗生活史策略关系在大尺度的群落空间水平上和小尺度的物种间与物种内的研究对于天然植被的更新恢复研究具有重要意义.     

Intrinsic biotic factors and microsite conditions drive seedling survival in a species with masting reproduction

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Quantitative genetic analysis of seed vigour and pre-emergence seedling growth traits in Brassica oleracea

The ability of seeds to germinate and establish seedlings in a predictable manner under a range of conditions has a direct contribution to the economic success of commercial crops, and should therefore be considered in crop improvement. We measured traits associated with seed vigour and pre-emergence seedling growth in a segregating population of 105 doubled haploid Brassica oleracea lines. The germination traits measured were: mean germination times for unstressed germination; germination under water stress or germination after a heat treatment; and conductivity of seed leachate. The seedling growth traits measured were: seed weight; seedling growth rate; and seedling size at the end of the exponential growth phase. There were some correlations, notably among germination traits, and between seed weight and pre-emergence seedling growth. Heritability of the various traits was typically in the 10–15% range, with heritability of conductivity and mean germination time under water stress 25 and 24% respectively. Collectively the results indicate that germination and pre-emergence seedling growth are under separate genetic control. Quantitative trait loci analyses were carried out on all measurements and revealed significant loci on linkage groups O1, O3, O6, O7 and O9. We suggest that genes at these loci are important in determining predictable seed germination and seedling establishment in practice.     

Timing of cotyledon damage affects growth and flowering in mature plants

Although the effects of herbivory on plant fitness are strongly linked to age, we understand little about how the timing of herbivory at the seedling stage affects growth and reproduction for plants that survive attack. In this study, we subjected six north-western European, dicotyledonous grassland species (Leontodon autumnalis, Leontodon hispidus, Plantago lanceolata, Plantago major, Trifolium pratense and Trifolium repens) to cotyledon removal at 7, 14 and 21 d old. We monitored subsequent growth and flowering (number of inflorescences recorded, and time taken for first flowers to open) over a 107 d period. Cotyledon removal reduced growth during establishment (35 d) for all species, and a further three exhibited reduced growth at maturity. Four species developed fewer inflorescences, or had delayed flowering after cotyledon removal. Although early damage (7 d old) had the greatest long-term effect on plant performance, responses varied according to the age at which the damage occurred and the species involved. Our results illustrate how growth and flowering into the mature phase is affected by cotyledon damage during different stages of seedling ontogeny, and we highlight the ways in which ontogenetic variation in seedling tolerance of tissue loss might impact upon plant fitness in mature plant communities.     

Equivalence in the strength of deer herbivory on above and below ground communities

Herbivores exert a strong influence on the species composition and richness of plant communities, but the magnitude of their effect on belowground communities remains poorly understood. While an increasing number of studies acknowledge the importance of documenting belowground effects of herbivores, very few of these evaluate variation in the strength of the response from aboveground to belowground communities. Our study documents the long-term consequences of sustained deer herbivory for plant and arthropod communities adjacent to 15 exclosures that have been in place since 1996. We hypothesized that herbivory would alter the composition and diversity of communities, but the strength of the effects of herbivory would weaken from plants, to leaf-litter invertebrates, and to belowground microarthropod communities. First, we found that herbivory negatively impacted plant seedling and sapling abundance and performance, reduced the abundance of ants and the taxonomic richness of arthropods in the litter layer and reduced the richness of soil microarthropod communities. Second, in contrast to our hypothesis, the magnitude of effect size did not vary among trophic levels, indicating that effects of deer herbivory cascade from plants to the leaf-litter and soil arthropod communities with equal strength. While much recent research has focused on how specific traits of plants may mediate the effects of herbivory on associated species, our results suggest that indirect effects of herbivory might influence many components of belowground communities.     

Climate change and plant regeneration from seed

At the core of plant regeneration, temperature and water supply are critical drivers for seed dormancy (initiation, break) and germination. Hence, global climate change is altering these environmental cues and will preclude, delay, or enhance regeneration from seeds, as already documented in some cases. Along with compromised seedling emergence and vigour, shifts in germination phenology will influence population dynamics, and thus, species composition and diversity of communities. Altered seed maturation (including consequences for dispersal) and seed mass will have ramifications on life history traits of plants. Predicted changes in temperature and precipitation, and thus in soil moisture, will affect many components of seed persistence in soil, e.g. seed longevity, dormancy release and germination, and soil pathogen activity. More/less equitable climate will alter geographic distribution for species, but restricted migratory capacity in some will greatly limit their response. Seed traits for weedy species could evolve relatively quickly to keep pace with climate change enhancing their negative environmental and economic impact. Thus, increased research in understudied ecosystems, on key issues related to seed ecology, and on evolution of seed traits in nonweedy species is needed to more fully comprehend and plan for plant responses to global warming.     

Plant recruitment and range size: Transition probabilities from ovule to adult in two Ipomoea

In plants, narrow geographic distributions are generally associated with low colonization and persistence abilities, therefore narrowly distributed plants are expected to have lower plant recruitment success than widespread species. Determining the association between recruitment success and range size requires the comparison of the success in multiple life-history stages among narrowly distributed and widespread congeners sharing the same habitat, an integrated approach rarely considered. We compared transition probabilities from ovule to reproductive adult between the narrowly distributed annual vine Ipomoea rubriflora O’Donnell (Convolvulaceae) and the widespread Ipomoea purpurea (L.) Roth at sites in Chaco woodland where they co-occur. I. rubriflora had marginally lower ovule fertilization success, a lower probability of seed maturation and lower seedling establishment than I. purpurea. The lowest transition probability for both species was seedling emergence. Seedling establishment in I. rubriflora was similar to seedling emergence. Plant recruitment success in I. rubriflora was an order of magnitude lower than that of I. purpurea. Indeed, I. rubriflora had lower total seed production per plant and smaller seed mass. Our results suggest that understanding processes determining regeneration (i.e., seedling emergence and establishment) may be of high importance for understanding narrow distributions in annual plants. The results also suggest that the narrowly distributed species is not only at a “numerical disadvantage” for colonizing new sites—i.e., lower total seed production per plant—but it also has a lower per-seed probability of establishing and thus to persist. However, as lower establishment success in this species seemed to be associated with the lower ability to survive adverse conditions of seedlings from smaller-seeded species, a narrower regeneration niche might also explain its narrow distribution.     

Herbivory differentially affects male and female reproductive traits of Cucumis sativus

Herbivory is an important selection pressure in the life history of plants. Most studies use seed or fruit production as an indication of plant fitness, but the impact of herbivory on male reproductive success is usually ignored. It is possible that plants compensate for resources lost to herbivory by shifting the allocation from seed production to pollen production and export, or vice versa. This study examined the impact of herbivory by Helix aspersa on both male and female reproductive traits of a monoecious plant, Cucumis sativus. The effects of herbivory on the relative allocation to male and female flowers were assessed through measurements of the number and size of flowers of both sexes, and the amount of pollinator visitation. We performed two glasshouse experiments; the first looked at the impact of three levels of pre-flowering herbivory, and the second looked at four levels of herbivory after the plants had started to flower. We found that herbivory during the flowering phase led to a significant increase in the number of plants without male flowers. As a consequence there was significantly less pollen export from this population, as estimated by movement of a pollen analog. The size of female flowers was reduced by severe herbivory, but there was no affect on pollen receipt by the female flowers of damaged plants. The decrease in allocation to male function after severe herbivory may be adaptive when male reproductive success is very unpredictable.     

Chronic herbivory negatively impacts cone and seed production,seed quality and seedling growth of susceptible pinyon pines

Although herbivory often reduces the reproduction of attacked trees, few studies have examined how naturally occurring insect-resistant and susceptible trees differ in their reproduction, nor have these effects been experimentally examined through long-term herbivore removals. In addition, few studies have examined the effects of herbivory on the quality of seeds produced and the implications of reduced seed quality on seedling establishment. We evaluated the impact of chronic herbivory by the stem-boring moth, Dioryctria albovittella, on cone and seed production of the pinyon pine (Pinus edulis) during two mast years. Three patterns emerged. First, moth herbivory was associated with reductions in cone production, viable seed production and seed mass. Specifically, pinyons susceptible to moth attack had 93–95% lower cone production, and surviving cones produced 31–37% fewer viable seeds, resulting in a 96–97% reduction in whole tree viable seed production. In addition, surviving seeds from susceptible trees had 18% lower mass than resistant trees. Second, long-term experimental removal of the herbivore resulted in increased rates of cone and seed production and quality, indicating that moth herbivory was the driver of these reductions. Third, seed size was positively associated with seed germination and seedling biomass and height, suggesting that trees suffering chronic herbivory produce poorer quality offspring. Thus, the resistance traits of pinyons can affect the quality of offspring, which in turn may affect subsequent seedling establishment and population dynamics.     

How do pollinator visitation rate and seed set relate to species’ floral traits and community context?

Differences among plant species in visitation rate and seed set within a community may be explained both by the species’ floral traits and the community context. Additionally, the importance of species’ floral traits vs. community context on visitation rate and seed set may vary among communities. In communities where the pollinator-to-flower ratio is low, floral traits may be more important than community context, as pollinators may have the opportunity to be choosier when visiting plant species. In this study we investigated whether species’ floral traits (flower shape, size and number, and flowering duration) and community context (conspecific and heterospecific flower density, and pollinator abundance) could explain among-species variation in visitation rate and seed set. For this, we used data on 47 plant species from two Norwegian plant communities differing in pollinator-to-flower ratio. Differences among species in visitation rate and seed set within a community could be explained by similar variables as those explaining visitation rate and seed set within species. As expected, we found floral traits to be more important than community context in the community with a lower pollinator-to-flower ratio; whereas in the community with a higher pollinator-to-flower ratio, community context played a bigger role. Our study gives significant insights into the relative importance of floral traits on species’ visitation rate and seed set, and contributes to our understanding of the role of the community context on the fitness of plant species.     

Seed size, seedling morphology, and response to deep shade and damage in neotropical rain forest trees

To investigate the existence of coordinated sets of seedling traits adapted to contrasting establishment conditions, we examined evolutionary convergence in seedling traits for 299 French Guianan woody plant species and the stress response in a shadehouse of species representing seed size gradients within five major cotyledon morphology types. The French Guianan woody plant community has larger seeds than other tropical forest communities and the largest proportion of hypogeal cotyledon type (59.2%) reported for tropical forests. Yet the community includes many species with intermediate size seeds that produce seedlings with different cotyledonal morphologies. A split-plot factorial design with two light levels (0.8% and 16.1% PAR) and four damage treatments (control, seed damage, leaf damage, stem damage) was used in the shadehouse experiment. Although larger-seeded species had higher survival and slower growth, these patterns were better explained by cotyledon type than by seed mass. Even larger-seeded species with foliar cotyledons grew faster than species with reserve-type cotyledons, and survival after stem grazing was five times higher in seedlings with hypogeal cotyledons than with epigeal cotyledons. Thus, to predict seedling performance using seed size, seedling morphology must also be considered.