Changes in Forest Soil Properties in Different Successional Stages in Lower Tropical China

Background

Natural forest succession often affects soil physical and chemical properties. Selected physical and chemical soil properties were studied in an old-growth forest across a forest successional series in Dinghushan Nature Reserve, Southern China.

Methodology/Principal Findings

The aim was to assess the effects of forest succession change on soil properties. Soil samples (0–20 cm depth) were collected from three forest types at different succession stages, namely pine (Pinus massoniana) forest (PMF), mixed pine and broadleaf forest (PBMF) and monsoon evergreen broadleaf forest (MEBF), representing early, middle and advanced successional stages respectively. The soil samples were analyzed for soil water storage (SWS), soil organic matter (SOM), soil microbial biomass carbon (SMBC), pH, NH₄ ⁺-N, available potassium (K), available phosphorus (P) and microelements (available copper (Cu), available zinc (Zn), available iron (Fe) and available boron (B)) between 1999 and 2009. The results showed that SWS, SOM, SMBC, Cu, Zn, Fe and B concentrations were higher in the advanced successional stage (MEBF stage). Conversely, P and pH were lower in the MEBF but higher in the PMF (early successional stage). pH, NH₄ ⁺-N, P and K declined while SOM, Zn, Cu, Fe and B increased with increasing forest age. Soil pH was lower than 4.5 in the three forest types, indicating that the surface soil was acidic, a stable trend in Dinghushan.

Conclusion/Significance

These findings demonstrated significant impacts of natural succession in an old-growth forest on the surface soil nutrient properties and organic matter. Changes in soil properties along the forest succession gradient may be a useful index for evaluating the successional stages of the subtropical forests. We caution that our inferences are drawn from a pseudo-replicated chronosequence, as true replicates were difficult to find. Further studies are needed to draw rigorous conclusions regarding on nutrient dynamics in different successional stages of forest.     

Increasing phosphorus limitation along three successional forests in southern China

Background and Aims

Phosphorus (P) is commonly one of most limiting nutrients in tropical and subtropical forests, but whether P limitation would be exacerbated during forest succession remains unclear.

Methods

Soil phosphatase activity is often used as an indicator of P limitation. Here we examined soil acid phosphatase activity (APA) underneath tree species in pine forest (PF), mixed pine and broadleaf forest (MF) and monsoon evergreen broadleaf forest (MEBF) which represented the early, middle and late successional stages of subtropical forests in China, respectively. We also analyzed other indicators of P status (soil available P and N and P stoichiometry of the tree species).

Results

APA or APA per unit soil organic carbon under tree species was relatively low in the early successional forest. Different from PF and MF, soil available P beneath the tree species was lower than in the bulk soils in MEBF. Soil APA was closely related to N:P ratios of tree species across all three forests.

Conclusions

Our results imply that P limitation increases during forest succession at our site. The dominant tree species with low soil APAs in MEBF are likely more P-limited than other tree species.     

Secondary succession and effects of clear-logging on diversity in the subtropical forests on Okinawa Island,southern Japan

In order to clarify the recovery process of the subtropical forest on Okinawa Island, southern Japan, biomass accumulation and the successional trend of species diversity with time were investigated by comparing plots of old-growth and clear logged secondary forests. Self-thinning was an important factor in the development of young secondary forests, and the small variance in tree size within a stand was related to the stand not being fully stratified after clear-cutting. A large variance of size structure in old secondary and old-growth forests implies re-initiation of the understorey. Additionally, the trajectory of stand development indicated that the subtropical forest quickly recovered aboveground biomass, which reached its upper limit at about 50 years after disturbance. However, there was a large distinction in species diversity between the secondary forests and old-growth forests. The diversity of forest floor plants did not recover well after being clear-cut. This indicates that management of the subtropical forest should not only take timber-oriented tree species into account, but also the biodiversity in ground flora. The secondary forests were characterized by Castanopsis sieboldii and Schima wallichii, and the monopolization of C. sieboldii through secondary succession had a negative influence on species diversity. Distylium racemosum dominated at the late development stage and was considered a long-lived competing species that reduced the dominance of C. sieboldii and facilitated the co-occurrence of understorey species. Light-demanding pioneer tree species such as S. wallichii that regenerated after logging should be replaced by competitive effects of climax species, and thus relayed floristic change might increase species diversity along secondary succession.Nomenclature: Hatushima and Amano (1994).     

Demographic Drivers of Aboveground Biomass Dynamics During Secondary Succession in Neotropical Dry and Wet Forests

The magnitude of the carbon sink in second-growth forests is expected to vary with successional biomass dynamics resulting from tree growth, recruitment, and mortality, and with the effects of climate on these dynamics. We compare aboveground biomass dynamics of dry and wet Neotropical forests, based on monitoring data gathered over 3–16 years in forests covering the first 25 years of succession. We estimated standing biomass, annual biomass change, and contributions of tree growth, recruitment, and mortality. We also evaluated tree species’ contributions to biomass dynamics. Absolute rates of biomass change were lower in dry forests, 2.3 and 1.9 Mg ha^?1 y^?1, after 5–15 and 15–25 years after abandonment, respectively, than in wet forests, with 4.7 and 6.1 Mg ha^?1 y^?1, in the same age classes. Biomass change was largely driven by tree growth, accounting for at least 48% of biomass change across forest types and age classes. Mortality also contributed strongly to biomass change in wet forests of 5–15 years, whereas its contribution became important later in succession in dry forests. Biomass dynamics tended to be dominated by fewer species in early-successional dry than wet forests, but dominance was strong in both forest types. Overall, our results indicate that biomass dynamics during succession are faster in Neotropical wet than dry forests, with high tree mortality earlier in succession in the wet forests. Long-term monitoring of second-growth tropical forest plots is crucial for improving estimates of annual biomass change, and for enhancing understanding of the underlying mechanisms and demographic drivers.     

Contrasting Regeneration Strategies in Climax and Long-Lived Pioneer Tree Species in a Subtropical Forest

1

This study investigated 15 coexisting dominant species in a humid subtropical evergreen broad-leaved forest in southwest China, consisting of long-lived pioneers and climax species occurring in natural and disturbed regimes. The authors hypothesized that there would be non-tradeoff scaling relationships between sprouting and seed size among species, with the aim of uncovering the ecological relationship between plant sprouting and seed characteristics in the two functional groups.

2

The sprouting variations of the species were initially examined using pairwise comparisons between natural and disturbed habitats within and across species and were noted to show a continuum in persistence niches across the forest dominants, which may underlie the maintenance of plant diversity. Second, a significantly positive, rather than tradeoff, relationship between sprout number and seed size across species within each of the two functional groups was observed, and an obvious elevational shift with a common slope among the two groups in their natural habitat was examined. The results indicate the following: 1) the relationship of seed size vs. sprouts in the natural habitat is more likely to be bet-hedging among species within a guild in a forest; 2) climax species tend to choose seeding rather than sprouting regeneration, and vice versa for the long-lived pioneers; and 3) the negative correlation between sprouting and seed dispersal under disturbed conditions may imply a tradeoff between dispersal and persistence in situ during the process of plant regeneration.

3

These findings may be of potential significance for urban greening using native species.     

海南岛霸王岭热带低地雨林植被恢复动态

热带次生林具有重要的物种保育和固碳功能, 然而高强度的干扰会导致次生林早期出现类似季雨林的阶段, 因而群落恢复速度和方向是当前热带生态学研究中最为关注的议题之一。该文以海南岛在刀耕火种弃耕地形成的不同演替阶段的次生林为研究对象, 比较森林不同恢复时间(12年、25年、55年)群落中的不同年龄(幼树、小树、成年树)个体与低地雨林老龄林的物种组成、多样性和群落结构差异, 探讨刀耕火种弃耕地恢复过程中的群落组配过程。首先, 在海南岛霸王岭林区内建立7个1 hm²(100 m × 100 m)的样地, 并调查样地内所有胸径≥ 1 cm的木本植物个体(不包括木质藤本)的种类、胸径大小和树高。无度量多维标定法(NMS)排序结果表明, 刀耕火种弃耕地恢复群落与老龄林的物种组成存在明显差异, 并且其物种组成差异随着径级增加而逐渐减小。刀耕火种弃耕地群落物种累积速度缓慢, 25年和55年恢复群落的种面积、种个体和种多度曲线无差异, 存在一个明显的停滞阶段。与物种组成相比, 群落结构恢复相对较迅速, 但仍没有形成老龄林阶段中的复杂结构。萌生个体在早期恢复群落中占有较高比例, 其个体密度和胸高断面积分别占总数的39.9%和55.9%, 但在恢复中后期迅速降低。刀耕火种弃耕地恢复群落中以先锋种和非先锋喜光种为主。虽然耐阴种随演替而逐渐增加, 但恢复中后期群落中的耐阴种重要值仅为老龄林的27.7%。这些结果表明, 虽然刀耕火种弃耕地恢复群落缓慢地逐渐接近最终恢复目标, 但仍然存在 很大的不确定性。刀耕火种弃耕地恢复过程中的异速恢复和停滞阶段需要纳入今后群落演替模型构建和森林固碳效益核甘共苦算中。     

Standing fine root mass and production in four Chinese subtropical forests along a succession and species diversity gradient

Background and aims

The influences of succession and species diversity on fine root production are not well known in forests. This study aimed to investigate: (i) whether fine root biomass and production increased with successional stage and increasing tree species diversity; (ii) how forest type affected seasonal variation and regrowth of fine roots.

Methods

Sequential coring and ingrowth core methods were used to measure fine root production in four Chinese subtropical forests differing in successional stages and species diversity.

Results

Fine root biomass increased from 262 g·m^?2 to 626 g·m^?2 with increasing successional stage and species diversity. A similar trend was also found for fine root production, which increased from 86 to 114 g·m^?2 yr ^?1 for Cunninghamia lanceolata plantation to 211–240 g·m^?2 yr ^?1 for Choerospondias axillaries forest when estimated with sequential coring data. Fine root production calculated using the ingrowth core data ranged from 186 g·m^?2 yr ^?1 for C. lanceolata plantation to 513 g·m^?2 yr ^?1 for Lithocarpus glaber – Cyclobalanopsis glauca forest.

Conclusions

Fine root biomass and production increased along a successional gradient and increasing tree species diversity in subtropical forests. Fine roots in forests with higher species diversity exhibited higher seasonal variation and regrowth rate.     

Removal of cattle accelerates tropical dry forest succession in Northwestern Mexico

Domestic livestock influence patterns of secondary succession across forest ecosystems. However, the effects of cattle on the regeneration of tropical dry forests (TDF) in Mexico are poorly understood, largely because it is difficult to locate forests that are not grazed by cattle or other livestock. We describe changes in forest composition and structure along a successional chronosequence of TDF stands with and without cattle (chronic grazing or exclusion from grazing for ~ 8 year). Forest stands were grouped into five successional stages, ranging from recently abandoned to mature forest, for a total of 2.7 ha of the sampled area. The absence of cattle increased woody plant (tree and shrub) density and species richness, particularly in mid-successional and mature forest stands. Species diversity and evenness were generally greater in sites where cattle were removed and cattle grazing in early successional stands reduced establishment and/or recruitment of new individuals and species. Removal of cattle from forest stands undergoing succession appears to facilitate a progressive and non-linear change of forest structure and compositional attributes associated with rapid recovery, while cattle browsing acts as a chronic disturbance factor that compromises the resilience and structural and functional integrity of the TDF in northwestern Mexico. These results are important for the conservation, management, and restoration of Neotropical dry forests.     

Spatial and temporal structure of diversity and demographic dynamics along a successional gradient of tropical forests in southern Brazil

1. Analysis of the structure, diversity, and demographic dynamics of tree assemblages in tropical forests is especially important in order to evaluate local and regional successional trajectories.
2. We conducted a long‐term study to investigate how the structure, species richness, and diversity of secondary tropical forests change over time. Trees (DBH ≥ 5 cm) in the Atlantic Forest of southern Brazil were sampled twice during a 10‐year period (2007 and 2017) in six stands (1 ha each) that varied in age from their last disturbance (25, 60, 75, 90, and more than 100 years). We compared forest structure (abundance and basal area), species richness, alpha diversity, demographic rates (mortality, recruitment, and loss or gain in basal area), species composition, spatial beta diversity, and temporal beta diversity (based on turnover and nestedness indices) among stand ages and study years.
3. Demographic rates recorded in a 10‐year interval indicate a rapid and dynamic process of species substitution and structural changes. Structural recovery occurred faster than beta diversity and species composition recovery. The successional gradient showed a pattern of species trade‐off over time, with less spatial dissimilarity and faster demographic rates in younger stands. As stands grow older, they show larger spatial turnover of species than younger stands, making them more stochastic in relation to species composition. Stands appear to split chronologically to some extent, but not across a straightforward linear axis, reflecting stochastic changes, providing evidence for the formation of a nonequilibrium community.
4. Policy implications. These results reiterate the complexity and variability in forest succession and serve as a reference for the evaluation and monitoring of local management and conservation actions and for defining regional strategies that consider the diversity of local successional trajectories to evaluate the effectiveness of restoration measures in secondary forests of the Atlantic Forest biome.

     

Phyllostomid bat assemblages in different successional stages of tropical rain forest in Chiapas,Mexico

Due to their role in seed dispersal, changes in the community of phyllostomid bats have direct consequences on ecological succession. The objective of this work was to document changes in the structure of bat assemblages among secondary successional stages of tropical rain forest in Chiapas, Mexico. Bats were mist-netted at ground level during 18 months in 10 sites belonging to 3 successional stages: four sites represented early succession (2–8 years of abandonment), four intermediate succession (10–20 years of abandonment), and two late succession (mature old-growth forest).We captured 1,179 phyllostomids comprising 29 species. Phyllostomid species richness was 17 (58% of all species) in the early stage, 18 (62%) in the intermediate stage and 24 (83%) in the late stage. The late successional mature forest possessed nine species that were exclusively found there, whereas early and intermediate successional stages contained only one exclusive species. Sturnira lilium, Artibeus lituratus, Carollia perpicillata, Artibeus jamaicensis and Glossophaga soricina represented 88% of all captured phyllostomid bats. Frugivores made up more than 90% of the species captured in early and intermediate successional stages and 84% in late successional forest. The Bray–Curtis index of dissimilarity showed a replacement of species through successional stages with the largest dissimilarity between early and late stages, followed by intermediate and late, and the lowest dissimilarity between early and intermediate stages. The number of gleaning insectivore species increased during succession. The carnivorous guild was exclusively found in the late stage (three species). We conclude that the late successional mature forest was the main reservoir for the gleaning insectivore and carnivore guilds; however, early and intermediate successional stages possessed a great diversity of species including many frugivores.     

海南岛热带低地雨林刀耕火种弃耕地自然恢复过程中的群落构建

研究群落构建机制是群落生态学的一个重要目标, 群落动态过程中的构建规律对于了解群落演替机理有重要的作用。该文以海南岛刀耕火种干扰后自然恢复的10 hm ²热带低地雨林为研究对象, 通过比较不同恢复阶段的次生林(15年、30年和60年)和老龄林在幼苗、幼树和成年树群落的物种组成, 揭示次生演替过程中的群落构建规律。研究结果表明, 老龄林中不同径级群落的物种多样性及不同径级间的物种相似度显著高于各恢复阶段的次生林, 但优势种在群落中的比例低于各恢复阶段的次生林。随着自然恢复过程的进行, 次生林群落物种组成与老龄林的相似性也逐渐增大, 支持演替平衡理论。所有恢复阶段样地中幼苗的个体、物种丰富度和基于多度涵盖估计量(ACE)都低于幼树和成年树群落, 幼苗层物种组成与幼树、成年树也有较大差异, 说明新增到幼苗群落可能是一个难于预测的过程。研究结果说明了确定过程和随机过程共同决定了次生演替的群落构建。     

Disrupted montane forest recovery hinders biodiversity conservation in the tropical Andes

Aim

Andean montane forests are biodiversity hotspots and large carbon stores and they provide numerous ecosystem services. Following land abandonment after centuries of forest clearing for agriculture in the Andes, there is an opportunity for forest recovery. Field-based studies show that forests do not always recover. However, large-scale and long-term knowledge of recovery dynamics of Andean forests remains scarce. This paper analyses tropical montane forest recovery trajectories over a 15-year time frame at the landscape and tropical Andean scale to inform restoration planning.

Methods

We first detect “potential recovery” as areas that have experienced a forest transition between 2000 and 2005. Then, we use Landsat time series analysis of the normalized difference water index (NDWI) to classify four “realized recovery” trajectories (“ongoing”, “arrested”, “disrupted” and “no recovery”) based on a sequential pattern of 5-yearly Z-score anomalies for 2005–2020. We compare these results against an analysis of change in tree cover to validate against other datasets.

Results

Across the tropical Andes, we detected a potential recovery area of 274 km² over the period. Despite increases in tree cover, most areas of the Andes remained in early successional states (10–25% tree cover), and NDWI levelled out after 5–10 years. Of all potential forest recovery areas, 22% showed “ongoing recovery”, 61% showed either “disrupted” or “arrested recovery”, and 17% showed “no recovery”. Our method captured forest recovery dynamics in a Peruvian arrested succession context and in landscape-scale tree-planting efforts in Ecuador.

Main conclusions

Forest recovery across the Andes is mostly disrupted, arrested or unsuccessful, with consequences for biodiversity recovery and provision of ecosystem services. Low-recovery areas identified in this study might be good candidates for active restoration interventions in this UN Decade on Restoration. Future studies could determine restoration strategies and priorities and suggest management strategies at a local planning scale across key regions in the biodiversity hotspot.     

The role of breeding system in community dynamics: Growth and mortality in forests of different successional stages

Plant sexual systems appear to play an important role in community assembly: Dioecious species are found to tend to have a higher propensity to colonize communities in early successional stages. Here, we test two demographic hypotheses to explain this pattern in temperate forests. First, we test demographic differences between hermaphrodite and dioecious species in stressful younger successional stages: Previous theory predicts that hermaphrodite seed production is more harmed in stressful environments than that of dioecious populations leading to an advantage for females of dioecious species. Second, in primary forest, we hypothesized that dioecious species would show demographic advantage over monomorphic ones. We used data from two temperate forest plots in Northeast China surveyed over 10 years to compare the rates of growth and mortality of tree species with contrasting breeding systems in both secondary and primary forests. We assessed the effect of breeding system on the growth‐mortality trade‐off, while controlling for other traits usually considered as correlates of growth and mortality rates. We show that in the secondary forest, dioecious species showed weak advantage in demographic rates compared with monomorphic species; dioecious species showed considerably both lower relative growth and mortality rates compared to the hermaphrodites in the primary forest over 10 years, consistent with a priori predictions. Hermaphrodites showed strong growth‐mortality trade‐offs across forest stages, even when possibly confounding factors had been accounted for. These results suggest that sexual system influences community succession and assembly by acting on the rates of growth and mortality, and the trade‐off between them. As vegetation develops, the demographic differences between breeding systems are much larger. Our results demonstrate the association between breeding system, succession, and community assembly and that this relationship is succession‐stage dependent. Our findings support the suggestion that the demographic advantage of dioecious species facilitates the coexistence of sexual systems in primary forest.     

Dispersal mode,shade tolerance,and phytogeographical affinity of tree species during secondary succession in tropical montane cloud forest

Secondary succession following land abandonment, represented by a chronosequence of 15 old fields (0–80 years old) and two old-growth forests, was studied in the tropical montane cloud forest region of Veracruz, Mexico. The objective was to determine successional trajectories in forest structure and species richness of trees ≥5 cm DBH, in terms of differences in seed dispersal mode, shade tolerance, and phytogeographical affinity. Data were analyzed using AIC model selection and logistic regressions. Mean and maximum canopy height reached values similar to old-growth forest at 35 and 80 years, respectively. Species richness and diversity values were reached earlier (15 and 25 years, respectively) while basal area and stem density tended to reach old-growth forest values within 80 years. Along the chronosequence, the proportion of species and individuals of wind-dispersed trees declined, that of bird dispersed small seeded trees remained constant, while that of gravity and animal dispersed large seeded trees increased; shade-intolerant species and individuals declined, while intermediate and shade-tolerant trees increased. Shade-tolerant canopy trees were rare during succession, even in the old-growth forest. Tropical tree species were more frequent than temperate ones throughout the chronosequence, but temperate tree individuals became canopy dominants at intermediate and old-growth forest stages.     

Temporal variation in the influence of forest succession on caterpillar communities: A long‐term study in a tropical dry forest

Forest succession can influence herbivore communities through changes in host availability, plant quality, microclimate, canopy structure complexity and predator abundance. It is not well known, however, if such influence is constant across years. Caterpillars have been reported to be particularly susceptible to changes in plant community composition across forest succession, as most species are specialists and rely on the presence of their hosts. Nevertheless, in the case of tropical dry forests, plant species have less defined successional boundaries than tropical wet forests, and hence herbivore communities should be able to persist across different successional stages. To test this prediction, caterpillar communities were surveyed during eight consecutive years in a tropical dry forest in four replicated successional stages in Chamela, Jalisco and Mexico. Lepidopteran species richness and diversity were equivalent in mature forests and early successional stages, but a distinctive caterpillar community was found for the recently abandoned pastures. Species composition tended to converge among all four successional stages during the span of eight years. Overall, our results highlight the importance of both primary and secondary forest for the conservation of caterpillar biodiversity at a landscape level. We also highlight the relevance of long‐term studies when assessing the influence of forest succession to account for across year variation in species interactions and climatic factors. Abstract in French is available with online material.     

Measuring the Conservation Value of Tropical Primary Forests: The Effect of Occasional Species on Estimates of Biodiversity Uniqueness

Background

Developing effective conservation plans for multi-functional landscapes requires an accurate knowledge of the relative conservation value of different land-uses. A growing number of tropical ecologists have evaluated conservation value using the number (or proportion) of species that are unique to primary or old-growth forests. However, estimates of the conservation value of modified land-uses may be inflated by the presence of occasional species (e.g. singletons and doubletons) that may be unable to exist as viable populations in isolation.

Methodology/Principal Findings

We use a unique 15-taxa dataset from a mixed-use forest landscape in the Brazilian Amazon to test the hypothesis that the removal of occasional species from sample data can increase estimates of the value of primary forest for biodiversity conservation.

Conclusions/Significance

Estimates of conservation value that are based on the proportion of species that are unique to tropical primary or old-growth forests are highly sensitive to decisions researchers make regarding the inclusion or exclusion of occasional species. By removing singletons from modified forest samples, and considering only those species known to occur in primary forest, we almost double estimates of the conservation value of tropical primary forests.     

The role of land-use history in driving successional pathways and its implications for the restoration of tropical forests

Secondary forests are increasingly important components of human-modified landscapes in the tropics. Successional pathways, however, can vary enormously across and within landscapes, with divergent regrowth rates, vegetation structure and species composition. While climatic and edaphic conditions drive variations across regions, land-use history plays a central role in driving alternative successional pathways within human-modified landscapes. How land use affects succession depends on its intensity, spatial extent, frequency, duration and management practices, and is mediated by a complex combination of mechanisms acting on different ecosystem components and at different spatial and temporal scales. We review the literature aiming to provide a comprehensive understanding of the mechanisms underlying the long-lasting effects of land use on tropical forest succession and to discuss its implications for forest restoration. We organize it following a framework based on the hierarchical model of succession and ecological filtering theory. This review shows that our knowledge is mostly derived from studies in Neotropical forests regenerating after abandonment of shifting cultivation or pasture systems. Vegetation is the ecological component assessed most often. Little is known regarding how the recovery of belowground processes and microbiota communities is affected by previous land-use history. In published studies, land-use history has been mostly characterized by type, without discrimination of intensity, extent, duration or frequency. We compile and discuss the metrics used to describe land-use history, aiming to facilitate future studies. The literature shows that (i) species availability to succession is affected by transformations in the landscape that affect dispersal, and by management practices and seed predation, which affect the composition and diversity of propagules on site. Once a species successfully reaches an abandoned field, its establishment and performance are dependent on resistance to management practices, tolerance to (modified) soil conditions, herbivory, competition with weeds and invasive species, and facilitation by remnant trees. (ii) Structural and compositional divergences at early stages of succession remain for decades, suggesting that early communities play an important role in governing further ecosystem functioning and processes during succession. Management interventions at early stages could help enhance recovery rates and manipulate successional pathways. (iii) The combination of local and landscape conditions defines the limitations to succession and therefore the potential for natural regeneration to restore ecosystem properties effectively. The knowledge summarized here could enable the identification of conditions in which natural regeneration could efficiently promote forest restoration, and where specific management practices are required to foster succession. Finally, characterization of the landscape context and previous land-use history is essential to understand the limitations to succession and therefore to define cost-effective restoration strategies. Advancing knowledge on these two aspects is key for finding generalizable relations that will increase the predictability of succession and the efficiency of forest restoration under different landscape contexts.     

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.     

Composition and Dynamics of Functional Groups of Trees During Tropical Forest Succession in Northeastern Costa Rica

We compared the functional type composition of trees ≥10 cm dbh in eight secondary forest monitoring plots with logged and unlogged mature forest plots in lowland wet forests of Northeastern Costa Rica. Five plant functional types were delimited based on diameter growth rates and canopy height of 293 tree species. Mature forests had significantly higher relative abundance of understory trees and slow-growing canopy/emergent trees, but lower relative abundance of fast-growing canopy/emergent trees than secondary forests. Fast-growing subcanopy and canopy trees reached peak densities early in succession. Density of fast-growing canopy/emergent trees increased during the first 20 yr of succession, whereas basal area continued to increase beyond 40 yr. We also assigned canopy tree species to one of three colonization groups, based on the presence of seedlings, saplings, and trees in four secondary forest plots. Among 93 species evaluated, 68 percent were classified as regenerating pioneers (both trees and regeneration present), whereas only 6 percent were classified as nonregenerating pioneers (trees only) and 26 percent as forest colonizers (regeneration only). Slow-growing trees composed 72 percent of the seedling and sapling regeneration for forest colonizers, whereas fast-growing trees composed 63 percent of the seedlings and saplings of regenerating pioneers. Tree stature and growth rates capture much of the functional variation that appears to drive successional dynamics. Results further suggest strong linkages between functional types defined based on adult height and growth rates of large trees and abundance of seedling and sapling regeneration during secondary succession.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp     

Functional Trait Strategies of Trees in Dry and Wet Tropical Forests Are Similar but Differ in Their Consequences for Succession

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.