African savannas are the last stronghold of diverse large-mammal communities, and a major focus of savanna ecology is to understand how these animals affect the relative abundance of trees and grasses. However, savannas support diverse plant life-forms, and human-induced changes in large-herbivore assemblages—declining wildlife populations and their displacement by livestock—may cause unexpected shifts in plant community composition. We investigated how herbivory affects the prevalence of lianas (woody vines) and their impact on trees in an East African savanna. Although scarce (<2% of tree canopy area) and defended by toxic latex, the dominant liana,
Lianas and other climbing plants are structural parasites of trees, generally reducing host tree survival, growth, and reproduction, yet their influences on the outcome of competition among tree species have remained largely unexplored. We propose that there are three distinct components to liana–tree interactions: The focal tree species vary strongly in liana prevalence, load, tolerance, and burden. Interspecific variation in tolerance is the largest contributor to interspecific variation in burden. Species rankings of per capita population growth rates under current liana infestation levels differ somewhat from rankings under liana‐free conditions, and differ strongly from rankings under uniformly high liana infestation. Thus, lianas alter competitive hierarchies to benefit tree species that are relatively tolerant of and/or resistant to lianas. Among the focal tree species, shade‐tolerance is positively correlated with liana tolerance and prevalence, but largely unrelated to load and burden, meaning shade‐tolerance does not predict which species are competitively disadvantaged by lianas. We describe a variety of mechanisms by which lianas may potentially increase or decrease niche differences among tree species, including interactions with spatial and temporal environmental niche partitioning, and potential differences among tree species in relative vulnerability to different liana species.
- NSF-PAR ID:
- 10370787
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- Journal of Ecology
- Volume:
- 107
- Issue:
- 3
- ISSN:
- 0022-0477
- Page Range / eLocation ID:
- p. 1469-1481
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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Cynanchum viminale (Apocynaceae), was eaten by 15 wild large-herbivore species and was consumed in bulk by native browsers during experimental cafeteria trials. In contrast, domesticated ungulates rarely ate lianas. When we experimentally excluded all large herbivores for periods of 8 to 17 y (simulating extirpation), liana abundance increased dramatically, with up to 75% of trees infested. Piecewise exclusion of different-sized herbivores revealed functional complementarity among size classes in suppressing lianas. Liana infestation reduced tree growth and reproduction, but herbivores quickly cleared lianas from trees after the removal of 18-y-old exclosure fences (simulating rewilding). A simple model of liana contagion showed that, without herbivores, the long-term equilibrium could be either endemic (liana–tree coexistence) or an all-liana alternative stable state. We conclude that ongoing declines of wild large-herbivore populations will disrupt the structure and functioning of many African savannas in ways that have received little attention and that may not be mitigated by replacing wildlife with livestock. -
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Abstract Despite their low contribution to forest carbon stocks, lianas (woody vines) play an important role in the carbon dynamics of tropical forests. As structural parasites, they hinder tree survival, growth and fecundity; hence, they negatively impact net ecosystem productivity and long‐term carbon sequestration.
Competition (for water and light) drives various forest processes and depends on the local abundance of resources over time. However, evaluating the relative role of resource availability on the interactions between lianas and trees from empirical observations is particularly challenging. Previous approaches have used labour‐intensive and ecosystem‐scale manipulation experiments, which are infeasible in most situations.
We propose to circumvent this challenge by evaluating the uncertainty of water and light capture processes of a process‐based vegetation model (ED2) including the liana growth form. We further developed the liana plant functional type in ED2 to mechanistically simulate water uptake and transport from roots to leaves, and start the model from prescribed initial conditions. We then used the PEcAn bioinformatics platform to constrain liana parameters and run uncertainty analyses.
Baseline runs successfully reproduced ecosystem gas exchange fluxes (gross primary productivity and latent heat) and forest structural features (leaf area index, aboveground biomass) in two sites (Barro Colorado Island, Panama and Paracou, French Guiana) characterized by different rainfall regimes and levels of liana abundance.
Model uncertainty analyses revealed that water limitation was the factor driving the competition between trees and lianas at the drier site (BCI), and during the relatively short dry season of the wetter site (Paracou). In young patches, light competition dominated in Paracou but alternated with water competition between the wet and the dry season on BCI according to the model simulations.
The modelling workflow also identified key liana traits (photosynthetic quantum efficiency, stomatal regulation parameters, allometric relationships) and processes (water use, respiration, climbing) driving the model uncertainty. They should be considered as priorities for future data acquisition and model development to improve predictions of the carbon dynamics of liana‐infested forests.
Synthesis . Competition for water plays a larger role in the interaction between lianas and trees than previously hypothesized, as demonstrated by simulations from a process‐based vegetation model. -
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Abstract Hydraulic traits are important for woody plant functioning and distribution. Associations among hydraulic traits, other leaf and stem traits, and species’ performance are relatively well understood for trees, but remain poorly studied for lianas. We evaluated the coordination among hydraulic efficiency (i.e., maximum hydraulic conductivity), hydraulic safety (i.e., cavitation resistance), a suite of eight morphological and physiological traits, and species’ abundances for saplings of 24 liana species and 27 tree species in wet tropical forests in Panama. Trees showed a strong trade‐off between hydraulic efficiency and hydraulic safety, whereas efficiency and safety were decoupled in lianas. Hydraulic efficiency was strongly and similarly correlated with acquisitive traits for lianas and trees (e.g., positively with gas exchange rates and negatively with wood density). Hydraulic safety, however, showed no correlations with other traits in lianas, but with several in trees (e.g., positively with leaf dry matter content and wood density and negatively with gas exchange rates), indicating that in lianas hydraulic efficiency is an anchor trait because it is correlated with many other traits, while in trees both efficiency and safety are anchor traits. Traits related to shade tolerance (e.g., low specific leaf area and high wood density) were associated with high local tree sapling abundance, but not with liana abundance. Our results suggest that different, yet unknown mechanisms determine hydraulic safety and local‐scale abundance for lianas compared to trees. For trees, the trade‐off between efficiency and safety will provide less possibilities for ecological strategies. For lianas, however, the uncoupling of efficiency and safety could allow them to have high hydraulic efficiency, and hence high growth rates, without compromising resistance to cavitation under drought, thus allowing them to thrive and outperform trees under drier conditions.
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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 Lianas are prevalent in Neotropical forests, where liana‐tree competition can be intense, resulting in reduced tree growth and survival. The ability of lianas to grow relative to trees during the dry season suggests that liana‐tree competition is also strongest in the dry season. If correct, the predicted intensification of the drying trend over large areas of the tropics in the future may therefore intensify liana‐tree competition resulting in a reduced carbon sink function of tropical forests. However, no study has established whether the liana effect on tree carbon accumulation is indeed stronger in the dry than in the wet season.
Using 6 years of data from a large‐scale liana removal experiment in Panama, we provide the first experimental test of whether liana effects on tree carbon accumulation differ between seasons. We monitored tree and liana diameter increments at the beginning of the dry and wet season each year to assess seasonal differences in forest‐level carbon accumulation between removal and control plots.
We found that median liana carbon accumulation was consistently higher in the dry (0.52 Mg C ha−1year−1) than the wet season (0.36 Mg C ha−1year−1) and significantly so in three of the years. Lianas reduced forest‐level median tree carbon accumulation more severely in the wet (1.45 Mg C ha−1year−1) than the dry (1.05 Mg C ha−1year−1) season in all years. However, the relative effect of lianas was similar between the seasons, with lianas reducing forest‐level tree carbon accumulation by 46.9% in the dry and 48.5% in the wet season.
Synthesis. Our results provide the first experimental demonstration that lianas do not have a stronger competitive effect on tree carbon accumulation during the dry season. Instead, lianas compete significantly with trees during both seasons, indicating a large negative effect of lianas on forest‐level tree biomass increment regardless of seasonal water stress. Longer dry seasons are unlikely to impact liana‐tree competition directly; however, the greater liana biomass increment during dry seasons may lead to further proliferation of liana biomass in tropical forests, with consequences for their ability to store and sequester carbon.
