Seed dispersal, seedling establishment and gap partitioning among tropical pioneer trees

1 We examined the abundance and distribution patterns of pioneer seeds in the soil seed bank, and of pioneer seedlings in 53 recently formed gaps, in a 50‐ha forest dynamics plot on Barro Colorado Island (BCI), Panama. The aim was to assess the importance of dispersal limitation (failure of seeds to arrive at all sites suitable for their germination) and establishment limitation (failure of seeds having reached a site to germinate successfully and establish as seedlings) in determining patterns of gap occupancy.
2 The abundance of seeds in the soil seed bank was strongly negatively correlated with seed size, but was not correlated with the abundance of reproductive‐sized adult trees in the plot. In contrast, the abundance of pioneer seedlings > 10 cm height in natural gaps was strongly correlated with adult abundance, but was not correlated with seed size.
3 Seedlings were non‐randomly distributed among gaps, but seedling abundance was not directly related to gap size, and there was no evidence of partitioning of the light environment of gaps by small seedlings. Large differences in growth and mortality rates among species were observed after 1 year, and this may result in the gap size partitioning previously found in saplings of the same species.
4 Seedlings of most species, particularly those with large seeds, were relatively more abundant than expected in gaps close to their conspecific adults. Proximity to reproductives, and by inference dispersal limitation, therefore exerts some effect on seedling distribution. None the less, large differences between seed and seedling abundances for some species, and low seedling occupancy rates in some gaps close to adult conspecifics, suggest that seedling emergence probabilities and species‐specific establishment requirements may also be important determinants of local abundance.     

西双版纳野芭蕉先锋群落优势种群的生态位动态

运用Ｌｅｖｉｎｓ（１９６８）和Ｐｉａｎｋａ（１９７３）提出的方法研究西双版纳小果野芭蕉先锋群落优势种群的生态位动态特征。结果表明,该群落中生态位宽度值列前五位的种群是：小果野芭蕉（１．６１５７）,木奶果,榕树,接骨木和苎麻;小果野芭蕉的生态位宽度值随着演替进行逐渐增大;群落中降小果野芭焦与其余物种间生态位重叠值普遍较大外,接骨木,木奶果、苎麻和榕树和它种植物之间具有较大幅度的生态位重叠;在三个演替     

Why equalising trade-offs aren't always neutral

Equalising trade-offs, such as seed mass vs. number, have been invoked to reconcile neutral theory with observed differences between species. This is an appealing explanation for the dramatic seed size variation seen within guilds of otherwise similar plants: under size-symmetric competition, where resource capture is proportional to mass, the outcome of competition should be insensitive to whether species produce many small seeds or few large ones. However, under this assumption, stochastic variation in seed rain leads to exclusion of all but the smallest-seeded species. Thus stochasticity in seed arrivals, a process that was previously thought to generate drift, instead results in deterministic competitive exclusion. A neutral outcome is possible under one special case of a more general equalising framework, where seed mass affects survival but not competition. Further exploration of the feasibility of neutral trade-offs is needed to understand the respective roles of neutrality and niche structure in community dynamics.     

Seed supply, recruitment, and assembly: quantifying relative seed and establishment limitation in a plant community context

There is a growing consensus that the relative constraints of seed limitation and establishment limitation in recruitment strongly influence abundance patterns in plant communities. Although these constraints have direct relevance to coexistence, most investigations utilize a seed addition approach that offers limited insight into these dynamics. Here we report the results of an assembly experiment with annual plant species from California grasslands to examine how propagule pool characteristics (dominant species abundance, functional diversity) influence establishment and seed limitation (density independence and density dependence across a gradient of seed supply) for each species, as well as how these constraints affect community diversity. Species were predominantly colimited by seed and establishment constraints, exhibiting saturating recruitment functions with increased seed supply. Consistent with competition-colonization trade-off predictions, recruitment constraints often depended on the degree of seed limitation of the competitive dominant, Brassica nigra; diversity was greatest in communities where Brassica was seed limited. Functional similarity within the propagule pool did not affect recruitment across a range of seed supply; likewise, functional diversity of the propagule pool was not related to community diversity. We conclude that seed limitation of the dominant species rather than niche similarity influences interspecific competition for safe sites and scales up to affect community-level diversity.     

Are plant populations seed‐limited? A review of seed sowing experiments

We define seed limitation to be an increase in population size following seed addition. Here, we briefly consider how theoretical models deal with seed limitation and how seed sowing experiments can be used to unravel the extent of seed limitation in natural systems. We review two types of seed addition experiments: seed augmentation studies where seeds are added to existing populations; and seed introductions where seeds are sown in unoccupied sites. Overall, approximately 50% of seed augmentation experiments show evidence of seed limitation. These studies show that seed limitation tends to occur more commonly in early successional habitats and in early successional species. Most of the studies have concentrated on simply categorising populations as seed- or microsite-limited, but we believe that seed sowing experiments could be used to reveal much more about community structure, and we discuss possible future directions. In 53% of introduction studies (where seeds were sown at sites from which the species was known to be absent) the introduced species was recorded in at least one of the experimental sites following sowing. However, of the subset of studies where both seedlings and adult plants were recorded, 64% of sites contained seedlings while only 23% contained adults. This implies that, for many species, conditions for establishment are more stringent than conditions for germination. The successful establishment of plants in unoccupied patches indicates the potential for immigration to enhance local diversity (the spatial mass effect). Few studies continued monitoring for long enough to determine whether or not self-sustaining populations were successfully established, and no study attempted to link introduction sites to a putative natural source of propagules, or considered the dynamics of the metapopulation as a whole.     

Do seeds sense each other? Testing for density‐dependent germination in desert perennial plants

The timing of seedling emergence may strongly affect fitness in competitive environments. Therefore, selection should favour mechanisms that allow sensing neighbours prior to emergence. We tested whether or not germination is affected by density and identity of neighbouring seeds or seedlings of desert perennial plants. Based on theory, we predicted that germination fractions are independent of neighbouring seeds, that germination is accelerated in dense interspecific neighbourhoods, and neighbour effects are caused by seedlings, not by seeds. We examined germination fraction and timing of four naturally coexisting sandy desert perennial species in low versus high seed densities in both intra- and interspecific neighbourhoods, and with and without removal of newly emerged seedlings. Neighbours accelerated germination independent of density and this pattern was apparently caused by the presence of early emerging seedlings. Germination fractions were lower in high seed densities even when neighbours did not germinate, indicating that seeds were able to sense each other prior to emergence. Early germination may be adaptive because fast emerging seedlings may gain a competitive edge over slow emerging ones. However, since seeds that did not germinate died, delayed germination may only be advantageous for mother plants when sib competition is intense. Another key finding was a competitive hierarchy with late successional species germinating faster and inhibiting germination of pioneer species. This indicates that successional processes may be directed as early as during germination.     

The shifting balance of facilitation and competition affects the outcome of intra- and interspecific interactions over the life history of California grassland annuals

Trait-based resource competition in plants, wherein more similar plants compete more strongly for resources, is a foundation of niche-based explanations for the maintenance of diversity in plant communities. Alternatively, neutral theory predicts that community diversity can be maintained despite equivalent resource requirements among species. We examined interactions at three life history stages (germination, survival, and juvenile-adult growth) for three native and three exotic California annual species in a glasshouse experiment. We varied plant density and species composition in small pots, with pots planted with either intraspecific seeds or in a three species mix of intra- and interspecific neighbors. We saw a range of facilitative, neutral, and competitive interactions that varied significantly by species, rather than by native or exotic status. There were more competitive interactions at the emergence and juvenile-adult growth stages and more facilitative interactions for survival. Consequently, the relative strength of competition in intraspecific versus mixed-species communities depended on whether we considered only the juvenile-adult growth stage or the entire life history of the interacting plants. Using traditional analysis of juvenile-adult growth only, all species showed negative density-dependent interactions for final biomass production. However, when the net effect of plant interactions from seed to adult was considered, which is a prediction of population growth, two native species ceased to show negative density dependence, and the difference between intraspecific and mixed-species competition was only significant for one exotic species. Results were consistent with predictions of neutral, rather than niche, theory for five of six species.     

Seeds in droppings of tropical fruit-eating birds: importance of considering seed composition

Summary The probability of interspecific seedling competition from bird droppings is high because fruit-eating birds, on average, deposit more than two species in a single dropping. Moreover, birds vary both in the number of plant species they deposit in a given dropping and in the seed composition of those droppings. In a preliminary experiment, I examined effects of interspecific seedling competition from seeds found in bird droppings. Certain plant species were competitively superior in pairwise growth experiments using six common second-growth species. The survival of certain shrubs depended on which species it was disseminated with in bird droppings. Rapid germination time may promote competitive superiority in some cases. Birds affect plant fitness not only because of their behavior following dispersal, but also because they deposit seeds in different densities and combinations.     

Seedling disease in an annual legume: consequences for seedling mortality, plant size, and population seed production

The effect of seed and seedling mortality on plant population dynamics depends on the degree to which the growth and reproduction of surviving individuals can compensate for the deaths that occur. To explore this issue, we sowed seeds of the annual Kummerowia stipulacea at three densities in sunken pots in the field, which contained either field soil, microwaved field soil, or microwaved field soil augmented with oospores of three Pythium species. High sowing density reduced seedling establishment and seedling size, but these effects were independent of the soil treatment. In the oospore-augmented soil, seed and seedling survival was low. The surviving plants were initially smaller but, at maturity, average plant size was greatest in the oospore-augmented soil, compared to the other treatments. Total population seed production was unaffected by soil treatment, suggesting that the effect of disease was limited to the seedling stage, with surviving plants released from intraspecific competition. To test the hypothesis that the surviving plants in the oospore-augmented soil were more disease-resistant, seeds from each of the sowing density-soil type treatments were sown in a growth chamber inoculation study. No evidence for selection for resistance was found. A second inoculation experiment revealed that oospore inoculum reduced plant numbers and mass regardless of whether field or microwaved soil was used, suggesting that results from the field experiment were not dependent on the use of microwaved soil. The findings of this study indicate that the ecological effects of disease on individual plants and on plant populations are not necessarily equivalent. Received: 13 January 1999 / Accepted: 21 September 1999     

Tropical Forests Are Non-Equilibrium Ecosystems Governed by Interspecific Competition Based on Universal 1/6 Niche Width

Tropical forests are mega-diverse ecosystems that display complex and non-equilibrium dynamics. However, theoretical approaches have largely focused on explaining steady-state behaviour and fitting snapshots of data. Here we show that local and niche interspecific competition can realistically and parsimoniously explain the observed non-equilibrium regime of permanent plots of nine tropical forests, in eight different countries. Our spatially-explicit model, besides predicting with accuracy the main biodiversity metrics for these plots, can also reproduce their dynamics. A central finding is that tropical tree species have a universal niche width of approximately 1/6 of the niche axis that echoes the observed widespread convergence in their functional traits enabling them to exploit similar resources and to coexist despite of having large niche overlap. This niche width yields an average ratio of 0.25 between interspecific and intraspecific competition that corresponds to an intermediate value between the extreme claims of the neutral model and the classical niche-based model of community assembly (where interspecific competition is dominant). In addition, our model can explain and yield observed spatial patterns that classical niche-based and neutral theories cannot.     

Predicting competition coefficients for plant mixtures: reciprocity, transitivity and correlations with life-history traits

There are few empirical or theoretical predictions of how per capita or per individual competition coefficients for pairs of plant species should relate to each other. In contrast, there are a considerable number of general hypotheses that predict competitive ability as a function of a range of ecological traits, together with a suite of increasingly sophisticated models for competitive interactions between plant species. We re-analyse a data set on competition between all pairwise combinations of seven species and show that competition coefficients relate strongly to differences between the maximum sizes, root allocation, emergence time and seed size of species. Regressions suggest that the best predictor of competition coefficients is the difference in the maximum size of species and that correlations of the other traits with the competition coefficients occur through effects on the maximum size. We also explore the patterns of association between coefficients across the competition matrix. We find significant evidence for coefficient reciprocity (inverse relationships between the interspecific coefficients for species pairs) and transitivity (numerically predictable hierarchies of competition between species) across competition matrices. These results therefore suggest simple null models for plant community structure when there is competition for resources.     

植物病原菌在森林动态中的作用

植物病原菌作为森林生态系统的重要组成成分及调控因子之一,在森林动态中扮演着重要的角色。植物病原菌通过侵染过程导致寄主植物的幼苗及成熟个体死亡、成熟个体的种子量降低或不实,或造成植物个体或群落中不同物种不同程度的病害,影响它们之间的营养竞争,从而导致群落结构、物种及个体数量的变化。感染散布前、后的种子和土壤种子库中的种子,以及由种子萌发产生的幼苗,它们的存活率降低,进而影响森林中的种子散布、幼苗更新与增补格局。在天然林中,先锋树种比顶极树种对病原菌更敏感,群落演替的早期阶段对病原菌比较敏感。植物病原菌主要通过密度依赖机制造成森林树种不同的死亡格局,从而参与森林的动态过程。     

Effectiveness of seed sowing techniques for sloped restoration sites

Practitioners are challenged with choosing among many potentially effective methods for sowing seed in ecological restoration projects to achieve sufficient native plant establishment. We tested the effectiveness of seed sowing techniques on moderate and steep slopes in a Mediterranean climate by measuring native seedling density immediately following germination, as well as plant density, recruitment success, and soil movement through the second growing season. We calculated cost effectiveness of different methods as the native plant density per dollar spent sowing seed. While all sowing techniques resulted in significant native establishment compared with unseeded controls, hydro seeding on moderate slopes was the most cost effective (native seedlings established per dollar spent). Although all steep‐sloped seeding techniques resulted in high densities of native species, all methods also resulted in significant soil loss. Shrubs preferred hand seeding followed by jute netting on steep slopes, while forbs reached greatest densities with hydro seeding on moderate slopes. Seedlings of species with heavy seeds were present in greater densities than species with lighter seeds in imprint sowing treatments. The “best” seed sowing technique varied depending on slope and metric of success (native density, species richness, shrub density, or forb density). Different combinations of slope, technique, and success metric resulted in significantly different project costs, which implies opportunities for savings given careful decision‐making relative to mitigation needs on heterogeneous landscapes. Evaluations of techniques for restoring slopes are limited, yet critical for expanding the area capable of being restored and the application of limited conservation funding.     

Seed size as an indicator of seed quality: a case study of Primula veris

Seed size is a widely accepted measure of seed quality, because many earlier studies have shown that large seeds have high seedling survival, growth and establishment. We tested whether ovule loss increases size of the remaining seeds and whether such size increase affects seedling establishment. We removed all except one flower from inflorescences of Primula veris L. (Primulaceae), a perennial hemicryptophyte herb, at a late stage of flowering. Flower removal (FR) increased seed size by 33% compared to the control plants. We then divided the seeds within each treatment to small, middle-sized and large seeds and carried out a sowing experiment in the field. Within each experimental group, seedling establishment was positively associated with seed size. However, despite size differences, seeds from the FR and control groups had the same seedling establishment probability. Seeds from FR plants had a higher seedling emergence in May than those from control plants, but the number of seedlings alive per sowing plot in the late summer was the same in both experimental groups. Increase in seed mass after partial FR thus did not enhance seedling performance, although seed size variation due to other causes was positively correlated with seedling establishment. Further studies are needed to show whether plastic changes of seed size are usually adaptive or not.     

Seed size as an indicator of seed quality: a case study of Primula veris

Seed size is a widely accepted measure of seed quality, because many earlier studies have shown that large seeds have high seedling survival, growth and establishment. We tested whether ovule loss increases size of the remaining seeds and whether such size increase affects seedling establishment. We removed all except one flower from inflorescences of Primula veris L. (Primulaceae), a perennial hemicryptophyte herb, at a late stage of flowering. Flower removal (FR) increased seed size by 33% compared to the control plants. We then divided the seeds within each treatment to small, middle-sized and large seeds and carried out a sowing experiment in the field. Within each experimental group, seedling establishment was positively associated with seed size. However, despite size differences, seeds from the FR and control groups had the same seedling establishment probability. Seeds from FR plants had a higher seedling emergence in May than those from control plants, but the number of seedlings alive per sowing plot in the late summer was the same in both experimental groups. Increase in seed mass after partial FR thus did not enhance seedling performance, although seed size variation due to other causes was positively correlated with seedling establishment. Further studies are needed to show whether plastic changes of seed size are usually adaptive or not.     

Seed to seedling transitions in successional habitats across a tropical landscape

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed‐dispersal limitation and seedling‐establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; Φ_i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (E_it) were higher for animal‐dispersed than wind‐dispersed species, and for non‐pioneer than pioneer species. Non‐pioneers and animal‐dispersed species had higher expected relative recruit abundance (ε_it) in primary forest (median of 4 seeds recruit^?1) whereas in secondary forest wind‐dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind‐dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (Φ_i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of Φ_i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); Φ_i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.     

Do fitness-equalizing tradeoffs lead to neutral communities?

Neutral theory in ecology is aimed at describing communities where species coexist due to similarities rather than the classically posited niche differences. It assumes that all individuals, regardless of species identity, are demographically equivalent. However, Hubbell suggested that neutral theory may describe even niche communities because tradeoffs equalize fitness across species which differ in their traits. In fact, tradeoffs can involve stabilization as well as fitness equalization, and stabilization involves different dynamics and can lead to different community patterns than neutral theory. Yet the important question remains if neutral theory provides a robust picture of all fitness-equalized communities, of which communities with demographic equivalence are one special case. Here, I examine Hubbell’s suggestion for a purely fitness-equalizing interspecific birth–death tradeoff, expanding neutral theory to a theory describing this broader class of fitness-equalized communities. In particular, I use a flexible framework allowing examination of the influence of speciation dynamics. I find that the scaling of speciation rates with birth and death rates, which is poorly known, has large impacts on community structure. In most cases, the departure from the predictions of current neutral models is substantial. This work suggests that demographic and speciation complexities present a challenge to the future development and use of neutral theory in ecology as null model. The framework presented here will provide a starting point for meeting that challenge, and may also be useful in the development of stochastic niche models with speciation dynamics.     

Does the seed size/number trade‐off model determine plant community structure? An assessment of the model mechanisms and their generality

This paper examines four key mechanisms of the seed size/number trade-off (SSNT) models to assess their relevance to a general understanding of plant community structure. Mechanism 1 is that large seeds have a greater probability of winning in competition against smaller seeds. I provide interspecific experimental evidence that there is a competitive hierarchy among seedlings based on seed size. Mechanism 2 is that a trade-off exists between the number and size of seeds produced for a given reproductive allocation. Negative correlations between seed size and number were found consistently across a range of species from a range of habitats, from published literature. Mechanism 3, that seedling-seedling competition is an important influence on species composition, was found to exist potentially in a range of environments, including annual-dominated, post-fire and gap-dynamic communities. However, there is little quantitative evidence available and this is likely to be a restrictive mechanism. Mechanism 4, that small seeds are superior colonists due to their greater number, was tested in a field experiment in a calcareous grassland community. No supporting evidence was found, suggesting that the SSNT is not an important determinant of structure in this community. Thus two of the four mechanisms can be considered to hold true generally, while the third mechanism may be valid in particular environments. The fourth mechanism did not apply in the community tested, but could be tested in a wider range of communities.     

Co-evolution of seed size and seed predation

Using the evolutionarily stable strategy (ESS) approach in a model for the co-evolution of seed size and seed predation, I show that seed size variation within individual plants is favoured if there is a trade-off in the predator's attack rate for different seed sizes. A single seed size is not evolutionarily stable because a predator that is optimally adapted to one particular seed size cannot prevent invasion by plants with a different seed size. The model generates the following predictions. The ESS consists of a continuous range of seed sizes. Small seeds tend to be attacked more frequently than big seeds. Plants with many resources and plants with low (frequency-independent) juvenile mortality have more variable seeds than plants with few resources and a high juvenile mortality. Seed size variation is higher in fluctuating populations regulated by seed predation alone than in stable populations (partially) regulated by seedling competition. Predator searching behaviour does not directly affect the ESS seed size range, but may have an indirect effect by affecting population stability or the significance of seedling competition as a population regulating mechanism. Moreover, seed size distributions are found to be more skewed in favour of small seeds if predation is spatially non-uniform than if predation is more even. Application of the model to systems of several co-evolving plant and predator species is discussed.     

Niche differentiation in rainforest ant communities across three continents

A central prediction of niche theory is that biotic communities are structured by niche differentiation arising from competition. To date, there have been numerous studies of niche differentiation in local ant communities, but little attention has been given to the macroecology of niche differentiation, including the extent to which particular biomes show distinctive patterns of niche structure across their global ranges. We investigated patterns of niche differentiation and competition in ant communities in tropical rainforests, using different baits reflecting the natural food spectrum. We examined the extent of temporal and dietary niche differentiation and spatial segregation of ant communities at five rainforest sites in the neotropics, paleotropics, and tropical Australia. Despite high niche overlap, we found significant dietary and temporal niche differentiation in every site. However, there was no spatial segregation among foraging ants at the community level, despite strong competition for preferred food resources. Although sucrose, melezitose, and dead insects attracted most ants, some species preferentially foraged on seeds, living insects, or bird feces. Moreover, most sites harbored more diurnal than nocturnal species. Overall niche differentiation was strongest in the least diverse site, possibly due to its lower number of rare species. Both temporal and dietary differentiation thus had strong effects on the ant assemblages, but their relative importance varied markedly among sites. Our analyses show that patterns of niche differentiation in ant communities are highly idiosyncratic even within a biome, such that a mechanistic understanding of the drivers of niche structure in ant communities remains elusive.