The maintenance of tree diversity has been explained by multiple mechanisms. One of the most thoroughly studied is conspecific negative density dependence, in which specialist plant enemies reduce survivorship of seeds, seedlings or saplings located near adult conspecifics. Although there is much support that conspecific negative density dependence occurs in temperate forests, only a subset of the species investigated thus far exhibit this recruitment pattern. It remains unclear what drives differential susceptibility to conspecifics among tree species. Previous investigators have considered shade tolerance and mycorrhizal type (arbuscular mycorrhizal vs. ectomycorrhizal association) as two traits that might explain differential susceptibility to conspecific negative density dependence. Here, we test whether these two plant traits predict susceptibility of tree saplings to conspecific negative density dependence in a temperate hardwood forest using three responses: spatial point patterns of saplings, sapling growth and sapling survival. Spatial patterns of saplings indicate that shade tolerant species are less sensitive to conspecifics than shade intolerant species, but show no differences based on mycorrhizal type. Conversely, shade tolerant saplings exhibit reduced growth, but not survival, when located in areas with high conspecific density. We interpret this finding in light of the conservative functional strategies of shade tolerant species, which typically have low leaf nitrogen levels and slower growth to divert resources to tissue defence against enemies. We found an effect of mycorrhizal type interacting with adult conspecific density, where arbuscular mycorrhizal species show a greater reduction in growth than ectomycorrhizal species in areas dense with conspecifics.
Ecologists often invoke interspecific facilitation to help explain positive biodiversity–ecosystem function relationships in plant communities, but seldom test how it occurs. One mechanism through which one species may facilitate another is by ameliorating abiotic stress. Physiological experiments show that a chronic excess of light can cause stress that depresses carbon assimilation. If shading by a plant's neighbours reduces light stress enough, it may facilitate that plant's growth. If light is instead most often a limiting factor for photosynthesis, shading may have an adverse, competitive effect. In a temperate tree diversity experiment, we measured stem growth rates and photosynthetic physiology in broadleaf trees across a gradient of light availability imposed by their neighbours. At the extremes, trees experienced nearly full sun (monoculture), or were shaded by nearby fast‐growing conifers (shaded biculture). Most species had slower growth rates with larger neighbours, implying a net competitive effect. On the other hand, the two most shade‐tolerant species ( We conclude that although large neighbours can cause light limitation in shade‐intolerant species, they can also increase growth through abiotic stress amelioration in shade‐tolerant species. Finally, in shade‐intolerant
- NSF-PAR ID:
- 10452302
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Journal of Ecology
- Volume:
- 109
- Issue:
- 5
- ISSN:
- 0022-0477
- Page Range / eLocation ID:
- p. 2000-2018
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Abstract Synthesis . We conclude that the shade tolerance level and the mycorrhizal type of temperate forest saplings may influence how their growth and survival respond to the adult conspecific trees in their neighbourhoods. -
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Abstract Forest encroachment into savannas is a widespread phenomenon, the rate of which may depend on soil conditions, species composition or changes in stand structure. As savanna specialist trees are replaced by generalist species, rates of stand development may increase. Because generalists can persist in forests, they are likely to grow more quickly and survive longer in dense stands, compared to savanna specialists. Furthermore, the faster growth rates of generalists may allow them to overtop and outcompete savanna specialists, causing rapid species turnover.
We measured growth and survival of 6,147 individuals of 112 species of savanna and generalist tree species over a period of 10 years in an ecological reserve in Assis, São Paulo State, Brazil. We modelled growth and mortality as a function of soil texture and nutrients, tree size, competitive neighbourhood, and membership in savanna or generalist (species which can persist in forests and savannas) functional groups.
Tree growth and survival was strongly influenced by competition, as estimated by the basal area of trees taller than a focal tree. At the stand level, savanna species are unable to contribute basal area growth in closed stands, while generalist species continue to increase in basal area even at high stand basal area. This phenomenon is driven by differences in growth and mortality. Generalists grew faster than savanna species, both in height and diameter. This difference in growth rates led to savanna species becoming suppressed more rapidly than generalists. When suppressed, savanna species were more than twice as likely to die than were generalists. Soils had inconsistent and mostly weak effects which were difficult to separate from gradients of stand structure.
Synthesis . We demonstrate that the presence of generalist trees accelerates the rates of basal area accumulation due to their greater growth rates and tolerance of shading. Generalists outcompete savanna trees by growing faster in the open and overtopping savanna specialists. Due to the slow growth and high mortality of savanna species in the shade, they are unable to form closed‐canopy stands. Accounting for differences among functional types and development of vegetation structure is critical for modelling forest encroachment. -
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Abstract Understanding the mechanisms that promote the coexistence of hundreds of species over small areas in tropical forest remains a challenge. Many tropical tree species are presumed to be functionally equivalent shade tolerant species but exist on a continuum of performance trade‐offs between survival in shade and the ability to quickly grow in sunlight. These trade‐offs can promote coexistence by reducing fitness differences.
Variation in plant functional traits related to resource acquisition is thought to predict variation in performance among species, perhaps explaining community assembly across habitats with gradients in resource availability. Many studies have found low predictive power, however, when linking trait measurements to species demographic rates.
Seedlings face different challenges recruiting on the forest floor and may exhibit different traits and/or performance trade‐offs than older individuals face in the eventual adult niche. Seed mass is the typical proxy for seedling success, but species also differ in cotyledon strategy (reserve vs. photosynthetic) or other leaf, stem and root traits. These can cause species with the same average seed mass to have divergent performance in the same habitat.
We combined long‐term studies of seedling dynamics with functional trait data collected at a standard life‐history stage in three diverse neotropical forests to ask whether variation in coordinated suites of traits predicts variation among species in demographic performance.
Across hundreds of species in Ecuador, Panama and Puerto Rico, we found seedlings displayed correlated suites of leaf, stem, and root traits, which strongly correlated with seed mass and cotyledon strategy. Variation among species in seedling functional traits, seed mass, and cotyledon strategy were strong predictors of trade‐offs in seedling growth and survival. These results underscore the importance of matching the ontogenetic stage of the trait measurement to the stage of demographic dynamics.
Our findings highlight the importance of cotyledon strategy in addition to seed mass as a key component of seed and seedling biology in tropical forests because of the contribution of carbon reserves in storage cotyledons to reducing mortality rates and explaining the growth‐survival trade‐off among species.
Synthesis : With strikingly consistent patterns across three tropical forests, we find strong evidence for the promise of functional traits to provide mechanistic links between seedling form and demographic performance. -
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Abstract Forest structure and diversity can regulate tree vulnerability to damage by insects and pathogens. Past work suggests that trees with diverse neighbours should experience less leaf herbivory and less damage from specialist herbivores and diseases, and that the effect of neighbourhood diversity should be strongest at small spatial scales.
In an early stage temperate tree diversity experiment, we monitored damage from leaf removing herbivores, specialist (gallers and leaf miners) herbivores, and two specialist fungal diseases (maple leaf anthracnose and cedar apple gall rust) over 3 years. The experimental design included treatments that varied independently in phylogenetic and functional diversity and we made our analyses across four spatial scales (1–16 m2).
Neighborhood diversity simultaneously increased leaf removal for some species, decreased it for others, and had no effect on yet others. Height apparency—the difference between a focal plant’s height and its neighbours’—was the best single predictor of leaf removal across species and spatial scales, but the strength and direction of its effect were also species‐specific.
Specialist pathogens and fungal foliar diseases showed signs of associational resistance and susceptibility. Oaks (
Quercus spp.) were more resistant to leaf miners and maples were more resistant to anthracnose when surrounded by diverse neighbours (associational resistance). In contrast, birches (Betula papyrifera ) were more susceptible to leaf miners and eastern red cedars (Juniperus virginiana ) were more susceptible to cedar apple gall rust (Gymnosporangium juniperi‐virginianae ) infection in diverse environments (associational susceptibility).Herbivore and pathogen damage was better predicted by community structure and diversity at small spatial scales (1 and 4 m2) than large scales (9 and 16 m2), suggesting a characteristic spatial scale for these biodiversity‐ecosystem functioning effects.
Synthesis. Humans control forest diversity through selective harvesting and planting in natural stands and plantations. Our experimental demonstration of the role of local community structure and diversity in suppressing some forms of pest and pathogen damage to trees suggests that forest management can be most effective when diversity is considered at small spatial scales and the underlying biology of particular pests, pathogens, and hosts is taken into account. Pictured here: the “galls” formed by cedar apple gall rust (Gymnosporangium juniperae‐virginiae) on eastern red cedar (Juniperus virginiana) in early spring release wind‐dispersed teleospores. Junipers showed associational susceptibility: greater susceptibility to gall rust with more diverse neighbours. -
Gomory, Dusan (Ed.)The expansion of woody plants into grasslands and old fields is often ascribed to fire suppression and heavy grazing, especially by domestic livestock. However, it is also recognized that nutrient availability and interspecific competition with grasses and other woody plants play a role in certain habitats. I examined potential factors causing range- and niche expansion by the eastern redcedar Juniperus virginiana , the most widespread conifer in the eastern United States, in multifactorial experiments in a greenhouse. Historical records suggest that the eastern redcedar is a pioneer forest species, and may be replaced as the forest increases in tree density due to shading. Another possible factor that affects its distribution may be nutrient availability, which is higher in old fields and other disturbed lands than in undisturbed habitats. In its historic range, eastern redcedars are particularly abundant on limestone outcrops, often termed ‘cedar barrens’. However, the higher abundance on limestone could be due to reduced interspecific competition rather than a preference for high pH substrates. I manipulated shade, fertilization, lime, and interspecific competition with a common dominant tree, the post oak Quercus stellata . In a separate experiment, I manipulated fire and grass competition. I measured growth rates (height and diameter) and above- and belowground biomass at the end of both experiments. I also measured total non-structural carbohydrates and nitrogen in these plants. Shade was the most important factor limiting the growth rates and biomass of eastern redcedars. I also found that there were significant declines in nitrogen and non-structural carbohydrates when shaded. These results are consistent with the notion that the eastern redcedar is a pioneer forest species, and that shade is the reason that these redcedars are replaced by other tree species. In the second experiment, I found that a single fire had a negative effect on young trees. There was no significant effect of competition with grass, perhaps because the competitive effect was shading by grasses and not nutrient depletion. Overall, the effects of shade were far more apparent than the effects of fire.more » « less
