Mechanistic models of community assembly state that biotic and abiotic filters constrain species establishment through selection on their functional traits. Predicting this assembly process is ...hampered because few studies directly incorporate environmental measurements and scale up from species to community level and because the functional traits' significance is environment dependent. We analyzed community assembly by measuring structure, environmental conditions, and species traits of secondary forests in a species-rich tropical system. We found, as hypothesized, that community structure shaped the local environment and that strong relationships existed between this environment and the traits of the most successful species of the regeneration communities. Path and multivariate analyses showed that temperature and leaf traits that regulate it were the most important factors of community differentiation. Comparisons between the trait composition of the forest's regeneration, juvenile, and adult communities showed a consistent community assembly pattern. These results allowed us to identify the major functional traits and environmental factors involved in the assembly of dry-forest communities and demonstrate that environmental filtering is a predictable and fundamental process of community assembly, even in a complex system such as a tropical forest.
Global plant trait studies have revealed fundamental trade-offs in plant resource economics. We evaluated such trait trade-offs during secondary succession in two species-rich tropical ecosystems ...that contrast in precipitation: dry deciduous and wet evergreen forests of Mexico. Species turnover with succession in dry forest largely relates to increasing water availability and in wet forest to decreasing light availability. We hypothesized that while functional trait trade-offs are similar in the two forest systems, the successful plant strategies in these communities will be different, as contrasting filters affect species turnover. Research was carried out in 15 dry secondary forest sites (5-63 years after abandonment) and in 17 wet secondary forest sites (<1-25 years after abandonment). We used 11 functional traits measured on 132 species to make species-trait PCA biplots for dry and wet forest and compare trait trade-offs. We evaluated whether multivariate plant strategies changed during succession, by calculating a 'Community-Weighted Mean' plant strategy, based on species scores on the first two PCA-axes. Trait spectra reflected two main trade-off axes that were similar for dry and wet forest species: acquisitive versus conservative species, and drought avoiding species versus evergreen species with large animal-dispersed seeds. These trait associations were consistent when accounting for evolutionary history. Successional changes in the most successful plant strategies reflected different functional trait spectra depending on the forest type. In dry forest the community changed from having drought avoiding strategies early in succession to increased abundance of evergreen strategies with larger seeds late in succession. In wet forest the community changed from species having mainly acquisitive strategies to those with more conservative strategies during succession. These strategy changes were explained by increasing water availability during dry forest succession and increasing light scarcity during wet forest succession. Although similar trait spectra were observed among dry and wet secondary forest species, the consequences for succession were different resulting from contrasting environmental filters.
We tested whether and how functional composition changes with succession in dry deciduous and wet evergreen forests of Mexico. We hypothesized that compositional changes during succession in dry ...forest were mainly determined by increasing water availability leading to community functional changes from conservative to acquisitive strategies, and in wet forest by decreasing light availability leading to changes from acquisitive to conservative strategies. Research was carried out in 15 dry secondary forest plots (5-63 years after abandonment) and 17 wet secondary forest plots (<1-25 years after abandonment). Community-level functional traits were represented by community-weighted means based on 11 functional traits measured on 132 species. Successional changes in functional composition are more marked in dry forest than in wet forest and largely characterized by different traits. During dry forest succession, conservative traits related to drought tolerance and drought avoidance decreased, as predicted. Unexpectedly acquisitive leaf traits also decreased, whereas seed size and dependence on biotic dispersal increased. In wet forest succession, functional composition changed from acquisitive to conservative leaf traits, suggesting light availability as the main driver of changes. Distinct suites of traits shape functional composition changes in dry and wet forest succession, responding to different environmental filters.
ABSTRACT
Old‐growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human‐modified ...tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio‐economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land‐use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio‐temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well‐preserved biodiversity‐rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales.
Significance Although forest succession has been approached as a predictable process, successional trajectories vary widely, even among nearby stands with similar environmental conditions and ...disturbance histories. We quantified predictability and uncertainty during tropical forest succession using dynamical models describing the interactions among stem density, basal area, and species density over time. We showed that the trajectories of these forest attributes were poorly predicted by stand age and varied significantly within and among sites. Our models reproduced the general successional trends observed, but high levels of noise were needed to increase model predictability. These levels of uncertainty call into question the premise that successional processes are consistent over space and time, and challenge the way ecologists view tropical forest regeneration.
Although forest succession has traditionally been approached as a deterministic process, successional trajectories of vegetation change vary widely, even among nearby stands with similar environmental conditions and disturbance histories. Here, we provide the first attempt, to our knowledge, to quantify predictability and uncertainty during succession based on the most extensive long-term datasets ever assembled for Neotropical forests. We develop a novel approach that integrates deterministic and stochastic components into different candidate models describing the dynamical interactions among three widely used and interrelated forest attributesâstem density, basal area, and species density. Within each of the seven study sites, successional trajectories were highly idiosyncratic, even when controlling for prior land use, environment, and initial conditions in these attributes. Plot factors were far more important than stand age in explaining successional trajectories. For each site, the best-fit model was able to capture the complete set of time series in certain attributes only when both the deterministic and stochastic components were set to similar magnitudes. Surprisingly, predictability of stem density, basal area, and species density did not show consistent trends across attributes, study sites, or land use history, and was independent of plot size and time series length. The model developed here represents the best approach, to date, for characterizing autogenic successional dynamics and demonstrates the low predictability of successional trajectories. These high levels of uncertainty suggest that the impacts of allogenic factors on rates of change during tropical forest succession are far more pervasive than previously thought, challenging the way ecologists view and investigate forest regeneration.
Vegetation and environment change mutually during secondary succession, yet the idiosyncrasies of the vegetation effect on the understorey environment are poorly understood. To test whether the ...successional understorey environment changes predictably and is shaped by the structure and seasonality of tropical dry forests, we estimated basal area and vegetation cover, and measured understorey temperature, light and moisture conditions, in 17 plots forming a 60-y chronosequence and a mature forest. Light and air and soil temperature decreased with time (75−15% of open-sky radiation, 31.7−29.3 °C, and +2.5 °C to −0.5 °C relative to ambient, respectively), whereas relative humidity increased (67−74%). Soil water availability increased with early-successional development (−45 to −1 kPa) but decreased afterwards (to −18 kPa). The first axis of a PCA of the rainy-season environment explained 60% of the variation and was strongly related to air temperature and relative humidity. During tropical dry-forest succession, such factors may be more important than light, the reduction in which is not extreme compared with taller and more vertically stratified wet forests. Seasonality significantly affected the successional environmental gradients, which were marked mainly during the wet season. Environmental heterogeneity was higher in the wet than in the dry season, and larger for resources (light and water) than for conditions (temperature and humidity). The wet-season increase in environmental heterogeneity potentially creates differential growing scenarios; the environmental harshness of the dry season would mostly challenge seedling survival.
The role of geological substrate in shaping plant community functional diversity remains poorly understood. Considering the involvement of leaves in the energy, water, and nutrient economics of ...plants, we hypothesized that leaves experience geology‐related filtering, which in turn shapes their functional attributes and community leaf functional diversity on different substrates. We studied tropical dry forest communities on limestone and siliciclastic phyllite‐derived soils, comparing their functional diversity and soil physico‐chemical properties. We predicted the most benign habitat (less severe filter) to be associated with higher leaf functional diversity and an acquisitive strategy prevalence, while the more stressful habitat should show conservative leaf traits and lower leaf functional diversity. We measured six traits in 31 common tree species (representing ~80% of community crown cover): leaf area, specific leaf area, leaf thickness, leaf dry matter content, petiole length, and leaf blade narrowness. Leaf functional diversity was assessed through the functional trait dispersion metric. Intraspecific functional variation was examined in 25 species shared between substrates. The limestone substrate was more fertile (higher phosphorous) with higher water retention, while phyllite had higher nitrogen and lower humidity. Principal component analysis segregated plots by substrate, with limestone plots being more clustered. Community leaf functional diversity was higher in the limestone forest. Most species examined showed inter‐substrate trait differences in at least one leaf functional trait. The two substrates constituted distinct growth environments, with the more benign substrate associated with higher community leaf functional diversity. The intraspecific analysis revealed the prevalence of acquisitive traits in the more benign and more conservative traits in the more stressful habitat. This study advances our understanding of the role of geological substrate as an environmental filter in tropical dry forests, influencing leaf functional responses and emphasizing the importance of intraspecific functional variation.
This study sheds light on how lithological substrates can shape community leaf functional diversity by creating environmental filters of varying intensities. It highlights the importance of studying intraspecific functional variation and provides an understanding of the complex link between soil and leaf functional traits.
Aim: Tropical forests account for a quarter of the global carbon storage and a third of the terrestrial productivity. Few studies have teased apart the relative importance of environmental factors ...and forest attributes for ecosystem functioning, especially for the tropics. This study aims to relate aboveground biomass (AGB) and biomass dynamics (i.e., net biomass productivity and its underlying demographic drivers: biomass recruitment, growth and mortality) to forest attributes (tree diversity, community-mean traits and stand basal area) and environmental conditions (water availability, soil fertility and disturbance). Location: Neotropics. Methods: We used data from 26 sites, 201 1-ha plots and >92,000 trees distributed across the Neotropics. We quantified for each site water availability and soil total exchangeable bases and for each plot three key community-weighted mean functional traits that are important for biomass stocks and productivity. We used structural equation models to test the hypothesis that all drivers have independent, positive effects on biomass stocks and dynamics. Results: Of the relationships analysed, vegetation attributes were more frequently associated significantly with biomass stocks and dynamics than environmental conditions (in 67 vs. 33% of the relationships). High climatic water availability increased biomass growth and stocks, light disturbance increased biomass growth, and soil bases had no effect. Rarefied tree species richness had consistent positive relationships with biomass stocks and dynamics, probably because of niche complementarity, but was not related to net biomass productivity. Community-mean traits were good predictors of biomass stocks and dynamics. Main conclusions: Water availability has a strong positive effect on biomass stocks and growth, and a future predicted increase in (atmospheric) drought might, therefore, potentially reduce carbon storage. Forest attributes, including species diversity and community-weighted mean traits, have independent and important relationships with AGB stocks, dynamics and ecosystem functioning, not only in relatively simple temperate systems, but also in structurally complex hyper-diverse tropical forests.
Rainforests are undergoing severe deforestation and fragmentation worldwide. A huge amount of small forest patches are being created, but their value in conserving biodiversity and forest structure ...is still controversial. Here, we demonstrate that in a species-rich and moderately-managed Mexican tropical landscape small rainforest patches (<100 ha) can be highly valuable for the conservation of tree diversity and forest structure. These patches showed diverse communities of native plants, including endangered species, and a new record for the country. Although the number of logged trees increased in smaller patches, patch size was a poor indicator of basal area, stem density, number of species, genera and families, and community evenness. Cumulative species-area curves indicated that all patches had a similar contribution to the regional species diversity. This idea also was supported by the fact that patches strongly differed in floristic composition (high β-diversity), independently of patch size. Thus, in agreement with the land-sharing approach, our findings support that small forest patches in moderately-managed landscapes should be included in conservation initiatives to maintain landscape heterogeneity, species diversity, and ecosystem services.
Land-use change modifies the spatial structure of tropical landscapes, shaping global biodiversity patterns. Yet, it remains unknown how key ecological processes, such as seed dispersal, can be ...affected by changes in landscape patterns, and whether such effects differ among regions with different climate and disturbance intensity. We assessed the effect of five landscape metrics (forest cover, matrix openness, forest edge density, forest fragmentation, and interpatch distance) on the seed rain recorded in two Mexican fragmented regions (20 forest sites per region): the more deforested, defaunated, and windy Los Tuxtlas rainforest (LTX), and the better-preserved Lacandona rainforest (LAC). We quantified the proportions of dispersed tree species and their seeds, separately evaluating wind- and animal-dispersed species. Our findings support the hypothesis that forest loss is more important than fragmentation per se, negatively impacting the seed rain in both regions. As expected, landscape patterns were comparatively more important for wind-dispersed seeds in LTX, probably because of stronger wind events in this region. Specifically, proportions of wind-dispersed seeds and species decreased with increasing edge density, suggesting that forest edges prevent dispersal of winddispersed species, which may occur if edges create physical barriers that limit wind flow. This pattern can also be caused by source limitation in landscapes with more forest edges, as tree mortality rates usually increase at forest edges. The wind-dispersed seed rain was also positively related to matrix openness, especially in LTX, where wind flow can be favored by the dominance of treeless anthropogenic matrices. Surprisingly, the proportion of animal-dispersed seeds in LTX was positively related to matrix openness and patch isolation, suggesting that seed dispersers in more deforested regions may be forced to concentrate in isolated patches and use the available habitat more intensively. Yet, as expected, patch isolation limited winddispersed seeds in LAC. Therefore, dispersal (and potentially regeneration) of wind-dispersed trees is favored in regions exposed to stronger wind events, especially in landscapes dominated by regularly shaped patches surrounded by open areas. Conversely, animal-dispersed seeds are primarily favored by increasing forest cover. Preventing forest loss is therefore critical to promote animal seed dispersal and forest recovery in human-modified rainforests.
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