Natalia Norden

Resilience of tropical rain forests: tree community reassembly in secondary forests

Understanding the recovery dynamics of ecosystems presents a major challenge in the human-impacted tropics. We tested whether secondary forests follow equilibrium or non-equilibrium dynamics by evaluating community reassembly over time, across different successional stages, and among multiple life stages. Based on long-term and static data from six 1-ha plots in NE Costa Rica, we show that secondary forests are undergoing reassembly of canopy tree and palm species composition through the successful recruitment of seedlings, saplings, and young trees of mature forest species. Such patterns were observed over time within sites and across successional stages. Floristic reassembly in secondary forests showed a clear convergence with mature forest community composition, supporting an equilibrium model. This resilience stems from three key factors co-occurring locally: high abundance of generalist species in the regional flora, high levels of seed dispersal, and local presence of old-growth forest remnants.

Above-ground biomass and productivity in a rain forest of eastern South America

Abstract: The dynamics of tropical forest woody plants was studied at the Nouragues Field Station, central French Guiana. Stem density, basal area, above-ground biomass and above-ground net primary productivity, including the contribution of litterfall, were estimated from two large permanent census plots of 12 and 10 ha, established on contrasting soil types, and censused twice, first in 1992?1994, then again in 2000?2002. Mean stem density was 512 stems ha?1 and basal area, 30m2 ha?1. Stem mortality rate ranged between 1.51% and 2.06% y?1. In both plots, stem density decreased over the study period. Using a correlation between wood density and wood hardness directly measured by a Pilodyn wood tester,we found that the mean wood densitywas 0.63 g cm?3, 12% smaller than the mean of wood density estimated from the literature values for the species occurring in our plot. Above-ground biomass ranged from 356 to 398Mgha?1 (oven-dry mass), and it increased over the census period. Leaf biomass was 6.47Mg ha?1. Our total estimate of aboveground net primary productivity was 8.81 MgC ha?1 y?1 (in carbon units), not accounting for loss to herbivory, branchfalls, or biogenic volatile organic compounds, whichmay altogether account for an additional 1MgC ha?1 y?1. Coarse wood productivity (stem growth plus recruitment) contributed to 4.16 MgC ha?1 y?1. Litterfall contributed to 4.65MgC ha?1 y?1 with 3.16 MgC ha?1 y?1 due to leaves, 1.10 MgC ha?1 y?1 to twigs, and 0.39MgC ha?1 y?1 to fruits and flowers. The increase in above-ground biomass for both trees and lianas is consistentwith the hypothesis of a shift in the functioning of Amazonian rain forests driven by environmental changes, although alternative hypotheses such as a recovery from past disturbances cannot be ruled out at our site, as suggested by the observed decrease in stem density. Key Words: above-ground biomass, carbon, French Guiana, net primary productivity, tropical forest

Successional dynamics in Neotropical forests are as uncertain as they are predictable

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 attributes—stem 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.

Multiple successional pathways in human-modified tropical landscapes : New insights from forest succession, forest fragmentation and landscape ecology research

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.

Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest

Negative density dependence (NDD) and environmental filtering (EF) shape community assembly, but their relative importance is poorly understood. Recent studies have shown that seedlings mortality risk is positively related to the phylogenetic relatedness of neighbours. However, natural enemies, whose depredations often cause NDD, respond to functional traits of hosts rather than phylogenetic relatedness per se. To understand the roles of NDD and EF in community assembly, we assessed the effects on seedling mortality of functional similarity, phylogenetic relatedness and stem density of neighbouring seedlings and adults in a species-rich tropical forest. Mortality risks increased for common species when their functional traits departed substantially from the neighbourhood mean, and for all species when surrounded by close relatives. This indicates that NDD affects community assembly more broadly than does EF, and leads to the tentative conclusion that natural enemies respond to phylogenetically correlated traits. Our results affirm the prominence of NDD in structuring species-rich communities.

A novel statistical method for classifying habitat generalists and specialists

We develop a novel statistical approach for classifying generalists and specialists in two distinct habitats. Using a multinomial model based on estimated species relative abundance in two habitats, our method minimizes bias due to differences in sampling intensities between two habitat types as well as bias due to insufficient sampling within each habitat. The method permits a robust statistical classification of habitat specialists and generalists, without excluding rare species a priori. Based on a user-defined specialization threshold, the model classifies species into one of four groups: (1) generalist; (2) habitat A specialist; (3) habitat B specialist; and (4) too rare to classify with confidence. We illustrate our multinomial classification method using two contrasting data sets: (1) bird abundance in woodland and heath habitats in southeastern Australia and (2) tree abundance in second-growth (SG) and old-growth (OG) rain forests in the Caribbean lowlands of northeastern Costa Rica. We evaluate the multinomial model in detail for the tree data set. Our results for birds were highly concordant with a previous nonstatistical classification, but our method classified a higher fraction (57.7%) of bird species with statistical confidence. Based on a conservative specialization threshold and adjustment for multiple comparisons, 64.4% of tree species in the full sample were too rare to classify with confidence. Among the species classified, OG specialists constituted the largest class (40.6%), followed by generalist tree species (36.7%) and SG specialists (22.7%). The multinomial model was more sensitive than indicator value analysis or abundance-based phi coefficient indices in detecting habitat specialists and also detects generalists statistically. Classification of specialists and generalists based on rarefied subsamples was highly consistent with classification based on the full sample, even for sampling percentages as low as 20%. Major advantages of the new method are (1) its ability to distinguish habitat generalists (species with no significant habitat affinity) from species that are simply too rare to classify and (2) applicability to a single representative sample or a single pooled set of representative samples from each of two habitat types. The method as currently developed can be applied to no more than two habitats at a time.

Demographic drivers of successional changes in phylogenetic structure across life‐history stages in plant communities

To gain insight into the ecological processes driving community reassembly in disturbed ecosystems, we assessed the phylogenetic dispersion of early- and late-successional tree species occurring in lowland forests of northeastern Costa Rica. Early-successional species were more closely related than expected by chance, whereas late-successional species tended to be less closely related than expected by chance. Then, we evaluated temporal changes in the phylogenetic structure of seedling and tree assemblages in four 1-ha plots of secondary forests in this region. We found an increase in the phylogenetic evenness among tree individuals over time in all secondary tree assemblages, indicating that relatedness among tree individuals decreases as succession unfolds. This pattern was jointly promoted by recruitment and mortality processes, suggesting that increasing evenness was caused by the replacement of individuals of early-successional species from closely related lineages by late-successional species belonging to a wider diversity of lineages. Based on species occurrence, however, tree community reassembly did not show any significant phylogenetic trend over time. These results suggest that shifts in species abundance over succession have a greater impact on the phylogenetic structure of the community than the turnover of species. Seedling assemblages showed higher phylogenetic evenness than tree assemblages, suggesting that propagule colonization is an important process driving phylogenetic changes in species composition throughout succession. Overall, our findings showed that the phylogenetic structure of these successional communities varies at two temporal scales. At short timescales, decreased dominance by early-successional species over succession leads to increased evenness among tree individuals. At longer timescales, colonization processes result in increased phylogenetic evenness in seedling communities compared to tree communities, forecasting increasing phylogenetic evenness among adult individuals at late-successional stages.

Mast Fruiting Is a Frequent Strategy in Woody Species of Eastern South America

Background It is thought that mast seeding is a rare reproductive strategy in the tropics, since tropical climates are less variable, and fruit consumers tend to be more generalist in these regions. However, previous tests of this hypothesis were based on only few tropical datasets, and none from tropical South America. Moreover, reproductive strategies have been quantified based on the coefficient of variation of interannual seed production, an index that potentially confounds masting and high interannual variability in seed production. Methodology/Principal Findings We developed a new approach to model the monthly variability in seed production for 28 tree species, and 20 liana species monitored during 5 years in a tropical forest of Central French Guiana. We found that 23% of the species showed a masting pattern, 54% an annual fruiting pattern, and 23% an irregular fruiting pattern. The majority of masting species were trees (8 out of 11), most of them animal-dispersed. The classification into reproductive strategies based on the coefficient of variation was inconsistent with our results in nearly half of the cases. Conclusions/Significance Our study is the first to clearly evidence the frequency of the masting strategy in a tropical forest community of Eastern South America. The commonness of the masting strategy in tropical plants may promote species coexistence through storage dynamics.

Temporal and spatial variability in seedling dynamics: a cross-site comparison in four lowland tropical forests

Spatialandtemporalvariationinseedlingdynamicswasassessedusingrecordsofcommunity-wideseedling demographycollectedwithidenticalmonitoringmethodsatfourtropicallowlandforestsinPanama,Malaysia,Ecuador and French Guiana for periods of between 3 and 10 y. At each site, the fates of between 8617 and 391 777 seedlings were followed through annual censuses of the 370-1008 1-m 2 seedling plots. Within-site spatial and inter-annual variation in density, recruitment, growth and mortality was compared with among-site variability using Bayesian hierarchical modelling to determine the generality of each sites patterns and potential for meaningful comparisons amongsites.TheMalaysianforest,whichexperiencescommunity-widemasting,wasthemostvariableinbothseedling density and recruitment. However, density varied year-to-year at all sites (CVamong years at site =8-43%), driven largely by high variability in recruitment rates (CV =40-117%). At all sites, recruitment was more variable than mortality (CV =5-64%) or growth (CV =12-51%). Increases in mortality rates lagged 1 y behind large recruitment events. Within-site spatial variation and inter-annual differences were greater than differences among site averages in all rates, emphasizing the value of long-term comparative studies when generalizing how spatial and temporal variation drive patterns of recruitment in tropical forests.

Environmental gradients and the evolution of successional habitat specialization: a test case with 14 Neotropical forest sites

Successional gradients are ubiquitous in nature, yet few studies have systematically examined the evolutionary origins of taxa that specialize at different successional stages. Here we quantify successional habitat specialization in Neotropical forest trees and evaluate its evolutionary lability along a precipitation gradient. Theoretically, successional habitat specialization should be more evolutionarily conserved in wet forests than in dry forests due to more extreme microenvironmental differentiation between early and late-successional stages in wet forest. We applied a robust multinomial classification model to samples of primary and secondary forest trees from 14 Neotropical lowland forest sites spanning a precipitation gradient from 788 to 4000 mm annual rainfall, identifying species that are old-growth specialists and secondary forest specialists in each site. We constructed phylogenies for the classified taxa at each site and for the entire set of classified taxa and tested whether successional habitat specialization is phylogenetically conserved. We further investigated differences in the functional traits of species specializing in secondary vs. old-growth forest along the precipitation gradient, expecting different trait associations with secondary forest specialists in wet vs. dry forests since water availability is more limiting in dry forests and light availability more limiting in wet forests. Successional habitat specialization is non-randomly distributed in the angiosperm phylogeny, with a tendency towards phylogenetic conservatism overall and a trend towards stronger conservatism in wet forests than in dry forests. However, the specialists come from all the major branches of the angiosperm phylogeny, and very few functional traits showed any consistent relationships with successional habitat specialization in either wet or dry forests. Synthesis. The niche conservatism evident in the habitat specialization of Neotropical trees suggests a role for radiation into different successional habitats in the evolution of species-rich genera, though the diversity of functional traits that lead to success in different successional habitats complicates analyses at the community scale. Examining the distribution of particular lineages with respect to successional gradients may provide more insight into the role of successional habitat specialization in the evolution of species-rich taxa.