Plant functional traits are thought to drive biomass production and biogeochemical cycling in tropical forests, but it remains unclear how nitrogen (N)‐fixing legumes influence the functional traits of neighbouring trees and forest‐wide biomass dynamics. Further, the degree to which effects of N‐fixers are density‐dependent and may depend on stem size and spatial scale remains largely unknown. Here, we examine 30 years of stem demography data for ~20,000 trees in a lowland tropical forest in Trinidad that span a wide range of functional traits thought to drive above‐ground biomass (AGB) dynamics. These forests show positive but decreasing long‐term net AGB accumulation resulting from constant average productivity but increasing mortality of non‐fixing trees over time. We find that high abundance of N‐fixing trees is associated with compositional shifts in non‐fixer functional traits that confer lower competitive performance and biomass accumulation. Across tree size classes, most interactions between N‐fixers and non‐fixers were negative, density‐dependent, and strongest at smaller spatial scales.
Nitrogen (N)‐fixing trees fulfil a unique and important biogeochemical role in forests through their ability to convert atmospheric N2gas to plant‐available N. Due to their high N fixation rates, it is often assumed that N‐fixing trees facilitate neighbouring trees and enhance forest growth. This assumption is supported by some local studies but contradicted by others, leaving the overall effect of N‐fixing trees on forest growth unresolved. Here we use the US Forest Service's Forest Inventory and Analysis database to evaluate the effects of N‐fixing trees on plot‐scale basal area change and individual‐scale neighbouring tree demography across the coterminous US. First we discuss the average trends. At the plot and individual scales, N‐fixing trees do not affect the relative growth rates of neighbouring trees, but they facilitate recruitment and inhibit survival rates, suggesting that they are drivers of tree turnover in the coterminous US. At the plot scale, N‐fixing trees facilitate the basal area change of non‐fixing neighbours. In addition to the average trends, there is wide variation in the effect of N‐fixing trees on forest growth, ranging from strong facilitation to strong inhibition. This variation does not show a clear geographical pattern, though it does vary with certain local factors. N‐fixing trees facilitate forest growth when they are likely to be less competitive: under high N deposition and high soil moisture or when neighbouring trees occupy different niches (e.g. high foliar C:N trees and non‐fixing trees).
- PAR ID:
- 10453524
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Journal of Ecology
- Volume:
- 109
- Issue:
- 2
- ISSN:
- 0022-0477
- Page Range / eLocation ID:
- p. 877-887
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
More Like this
-
Abstract Synthesis . Overall, our findings suggest that local trait‐based interactions between N‐fixing and non‐fixing trees can influence long‐term carbon accumulation in tropical forests. -
Abstract Tree death due to lightning influences tropical forest carbon cycling and tree community dynamics. However, the distribution of lightning damage among trees in forests remains poorly understood.
We developed models to predict direct and secondary lightning damage to trees based on tree size, crown exposure and local forest structure. We parameterized these models using data on the locations of lightning strikes and censuses of tree damage in strike zones, combined with drone‐based maps of tree crowns and censuses of all trees within a 50‐ha forest dynamics plot on Barro Colorado Island, Panama.
The likelihood of a direct strike to a tree increased with larger exposed crown area and higher relative canopy position (emergent > canopy >>> subcanopy), whereas the likelihood of secondary lightning damage increased with tree diameter and proximity to neighbouring trees. The predicted frequency of lightning damage in this mature forest was greater for tree species with larger average diameters.
These patterns suggest that lightning influences forest structure and the global carbon budget by non‐randomly damaging large trees. Moreover, these models provide a framework for investigating the ecological and evolutionary consequences of lightning disturbance in tropical forests.
Synthesis . Our findings indicate that the distribution of lightning damage is stochastic at large spatial grain and relatively deterministic at smaller spatial grain (<15 m). Lightning is more likely to directly strike taller trees with large crowns and secondarily damage large neighbouring trees that are closest to the directly struck tree. The results provide a framework for understanding how lightning can affect forest structure, forest dynamics and carbon cycling. The resulting lightning risk model will facilitate informed investigations into the effects of lightning in tropical forests. -
Abstract Most forests are recovering from human land use, making it critical to understand the effect of disturbance on forest recovery. Forests of the eastern United States have a long history of land use, but it is unknown whether historical disturbances have contributed to their transition from ectomycorrhizal (ECM) to arbuscular mycorrhizal (AM) tree dominance. Disturbance may promote nitrogen (N)‐fixing trees in early succession, which can elevate soil N availability even after they die. Higher soil N availability may facilitate the competitive success of AM trees over ECM trees, but such ‘N fixer founder effects’ have not been empirically tested.
Here, we analysed data from three land‐use disturbances in a temperate forest historically dominated by ECM trees: selective‐cutting (ranging from 0 to 52 m2 ha−1), clear‐cutting and agricultural abandonment. These disturbances occurred at different times, but long‐term data capture 3–7 decades of forest recovery.
We found that the AM tree fraction in contemporary forests was 2, 4, and 6‐fold higher following selective‐cutting, clear‐cutting and agricultural abandonment, respectively, compared to forest composition in 1934. Across these disturbances we also observed an increasing abundance of the N fixer black locust immediately following disturbance. Using a simulation model parameterized by data from black locust, we estimated historical rates of symbiotic N fixation to understand the relationship between N fixation and AM dominance in individual plots. We found that N fixation was positively associated with the growth of ECM trees generally, and oak and hickory specifically, only following light selective‐cutting (<12 or <18 m2 ha−1basal area extraction, respectively). Following higher levels of selective‐cutting and clear‐cutting, N fixation was positively associated with the growth of AM trees, particularly red maple and tulip poplar. Agricultural abandonment led to AM dominance regardless of N fixation rates.
Synthesis and applications . Our findings suggest that common land use practices and black locust, a native N fixer, can reduce the dominance of ECM trees. If N fixers are likely to proliferate following disturbance, we might maintain ECM dominance by cutting trees at low densities and by applying prescribed fire to remove N. -
Abstract Carbon uptake by the terrestrial biosphere depends on supplies of new nitrogen (N) from symbiotic N fixation, but we lack a framework for scaling fixation accurately and for predicting its response to global change.
We scaled symbiotic N fixation from individual N fixers (i.e. plants that host N‐fixing bacteria), by quantifying three key parameters—the abundance of N fixers, whether they are fixing N and their N fixation rates. We apply this framework to black locust, a widespread N‐fixing tree in temperate forests of the eastern United States, and harness long‐term data from southern Appalachian forests to scale fixation from trees to the landscape and over succession.
Symbiotic N fixation at the landscape scale peaked in the first decade following forest disturbance, and then declined. This pattern was due to the declining density and declining fixation rates of individual black locust trees over succession. Independent of forest succession, and coincident with chronic atmospheric N deposition, we have evidence suggesting that nodule biomass produced by black locust trees has declined by 83% over the last three decades. This difference in nodule biomass translates to a maximum fixation rate of 11 kg N ha−1 year−1and a landscape average of 1.5 kg N ha−1 year−1in contemporary forests.
Synthesis . We find key controls on symbiotic N fixation by black locust over space and time, suggesting lower fixation rates in eastern deciduous forests than previous estimates. Our scaling framework can be applied to other N fixers to aid predictions of symbiotic N fixation and ecosystem response to global change. -
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.