Modeling the Complex Impacts of Timber Harvests to Find Optimal Management Regimes for Amazon Tidal Floodplain Forests

At the Amazon estuary, the oldest logging frontier in the Amazon, no studies have comprehensively explored the potential long-term population and yield consequences of multiple timber harvests over time. Matrix population modeling is one way to simulate long-term impacts of tree harvests, but this approach has often ignored common impacts of tree harvests including incidental damage, changes in post-harvest demography, shifts in the distribution of merchantable trees, and shifts in stand composition. We designed a matrix-based forest management model that incorporates these harvest-related impacts so resulting simulations reflect forest stand dynamics under repeated timber harvests as well as the realities of local smallholder timber management systems. Using a wide range of values for management criteria (e.g., length of cutting cycle, minimum cut diameter), we projected the long-term population dynamics and yields of hundreds of timber management regimes in the Amazon estuary, where small-scale, unmechanized logging is an important economic activity. These results were then compared to find optimal stand-level and species-specific sustainable timber management (STM) regimes using a set of timber yield and population growth indicators. Prospects for STM in Amazonian tidal floodplain forests are better than for many other tropical forests. However, generally high stock recovery rates between harvests are due to the comparatively high projected mean annualized yields from fast-growing species that effectively counterbalance the projected yield declines from other species. For Amazonian tidal floodplain forests, national management guidelines provide neither the highest yields nor the highest sustained population growth for species under management. Our research shows that management guidelines specific to a region’s ecological settings can be further refined to consider differences in species demographic responses to repeated harvests. In principle, such fine-tuned management guidelines could make management more attractive, thus bridging the currently prevalent gap between tropical timber management practice and regulation.     

Does selective logging stress tropical forest invertebrates? Using fat stores to examine sublethal responses in dung beetles

The increased global demand for tropical timber has driven vast expanses of tropical forests to be selectively logged worldwide. While logging impacts on wildlife are predicted to change species distribution and abundance, the underlying physiological responses are poorly understood. Although there is a growing consensus that selective logging impacts on natural populations start with individual stress‐induced sublethal responses, this literature is dominated by investigations conducted with vertebrates from temperate zones. Moreover, the sublethal effects of human‐induced forest disturbance on tropical invertebrates have never been examined. To help address this knowledge gap, we examined the body fat content and relative abundance of three dung beetle species (Coleoptera: Scarabaeinae) with minimum abundance of 40 individuals within each examined treatment level. These were sampled across 34 plots in a before‐after control‐impact design (BACI) in a timber concession area of the Brazilian Amazon. For the first time, we present evidence of logging‐induced physiological stress responses in tropical invertebrates. Selective logging increased the individual levels of fat storage and reduced the relative abundance of two dung beetle species. Given this qualitative similarity, we support the measurement of body fat content as reliable biomarker to assess stress‐induced sublethal effects on dung beetles. Understanding how environmental modification impacts the wildlife has never been more important. Our novel approach provides new insights into the mechanisms through which forest disturbances impose population‐level impacts on tropical invertebrates.     

The persistence and conservation of Borneo’s mammals in lowland rain forests managed for timber: observations, overviews and opportunities

Lowland rainforests on Borneo are being degraded and lost at an alarming rate. Studies on mammals report species responding in various ways to habitat changes that occur in commercial forestry concessions. Here we draw together information on the relationship between the ecological, evolutionary, and biogeographic characteristics of selected Bornean non-volant mammals, and their response to timber harvesting and related impacts. Only a minority of species show markedly reduced densities after timber harvesting. Nonetheless there are many grounds for concern as various processes can, and often do, reduce the viability of wildlife populations. Our review of what we know, and of current understanding, helps predict mammalian dynamics and subsequent mammal-induced ecosystem changes in logged forests. We identify groups of mammal species that, although largely unstudied, are unlikely to tolerate the impacts associated with timber harvesting. On a positive note we find and suggest many relatively simple and low-cost ways in which concession management practices might be modified so as to improve the value of managed forests for wildlife conservation. Improving forest management can play a vital role in maintaining the rich biodiversity of Borneo’s tropical rain forests.     

Molecular database for classifying Shorea species (Dipterocarpaceae) and techniques for checking the legitimacy of timber and wood products

The extent of tropical forest has been declining, due to over-exploitation and illegal logging activities. Large quantities of unlawfully extracted timber and other wood products have been exported, mainly to developed countries. As part of the export monitoring effort, we have developed methods for extracting and analyzing DNA from wood products, such as veneers and sawn timbers made from dipterocarps, in order to identify the species from which they originated. We have also developed a chloroplast DNA database for classifying Shorea species, which are both ecologically and commercially important canopy tree species in the forests of Southeast Asia. We are able to determine the candidate species of wood samples, based on DNA sequences and anatomical data. The methods for analyzing DNA from dipterocarp wood products may have strong deterrent effects on international trade of illegitimate dipterocarp products. However, the method for analyzing DNA from wood is not perfect for all wood products and need for more improvement, especially for plywood sample. Consequently, there may be benefits for the conservation of tropical forests in Southeast Asia.     

Variation in tropical forest growth rates: combined effects of functional group composition and resource availability

Rates of tree growth in tropical forests reflect variation in life history strategies, contribute to the determination of species' distributional limits, set limits to timber harvesting and control the carbon balance of the stands. Here, we review the resources that limit tree growth at different temporal and spatial scales, and the different growth rates and responses of functional groups defined on the basis of regeneration strategy, maximum size, and species' associations with particular edaphic and climatic conditions.Variation in soil water availability determines intra- and inter-annual patterns of growth within seasonal forests, whereas irradiance may have a more important role in aseasonal forests. Nutrient supply limits growth rates in montane forests and may determine spatial variation in growth of individual species in lowland forests. However, its role in determining spatial variation in stand-level growth rates is unclear. In terms of growth rate, we propose a functional classification of tropical tree species which contrasts inherently fast-growing, responsive species (pioneer, large-statured species), from slow-growing species that are less responsive to increasing resource availability (shade-bearers, small-statured species). In a semi-deciduous forest in Ghana, pioneers associated with high-rainfall forests with less fertile soils, had significantly lower growth rates than pioneers that are more abundant in low-rainfall forests with more fertile soils. These results match patterns found in seedling trials and suggest for pioneers that species' associations with particular environmental conditions are useful indicators of maximum growth rate.The effects of variation in resource availability and of inherent differences between species on stand-level patterns of growth will not be independent if the functional group composition of tropical forests varies along resource gradients. We find that there is increasing evidence of such spatial shifts at both small and large scales in tropical forests. Quantifying these gradients is important for understanding spatial patterns in forest growth rates.     

Prospects for conserving biodiversity in Amazonian extractive reserves

Non‐timber forest product (NTFP) extraction is a popular alternative to timber extraction that figures prominently in efforts to utilize tropical forests sustainably. But the ability to conserve biodiversity through NTFP management, particularly in extractive reserves in Amazonia, has remained untested. We found that intensive management of Euterpe oleracea (Palmae) fruit, one of the most important extractive products in the Amazon, has substantial impacts on biodiversity, whereas moderate management does not. We mimicked traditional levels of fruit harvest in a replicated experiment over one fruiting season. High‐intensity harvest (75% of fruits removed) reduced avian frugivore species diversity by 22%. Low‐intensity harvest (40% of fruits removed), however, had no effect on diversity. On a larger scale, we found that forests with enriched densities of E. oleracea supported more fruit‐eating birds but fewer non fruit‐eating birds than non‐enriched forests. Taken together, these results suggest that intensive NTFP management to meet market demands may trigger substantial ecological impacts, at least at the level of our study. E. oleracea harvest should be limited where conservation of biodiversity is a goal.     

The Effect of Logging on the Ground-Foraging Ant Community in Eastern Amazonia

In order to determine the suitability of ants as indicator organisms for habitat disruption in tropical forests, we studied the effects of both high and low impact logging on ant communities in northeastern Pará State, in the Brazilian Amazon. We collected ants from logged forests and unlogged forest sites with Winkler bags throughout the 1998 rainy season (January and April) and the following dry season (July and September). Both methods of timber harvesting showed impacts on ant community composition when compared with unlogged forest, although these impacts did not include modifications in total species richness or the relative contribution of each ant subfamily to the total number of species. Instead, logging induced alterations took place at the level of species and genera. A 2-fold reduction in the dominance of ants of the highly diverse genus Pheidole was associated with forest alterations in high-impact logging sites. Thus, logging in Amazonia can be seen to promote species shifts in ant communities, without, however, altering species richness. Ants of the genus Pheidole are potentially useful indicators for forest disturbances resulting from timber extraction.     

Trade‐offs between carbon stocks and timber recovery in tropical forests are mediated by logging intensity

Forest degradation accounts for ~70% of total carbon losses from tropical forests. Substantial emissions are from selective logging, a land‐use activity that decreases forest carbon density. To maintain carbon values in selectively logged forests, climate change mitigation policies and government agencies promote the adoption of reduced‐impact logging (RIL) practices. However, whether RIL will maintain both carbon and timber values in managed tropical forests over time remains uncertain. In this study, we quantify the recovery of timber stocks and aboveground carbon at an experimental site where forests were subjected to different intensities of RIL (4, 8, and 16 trees/ha). Our census data span 20 years postlogging and 17 years after the liberation of future crop trees from competition in a tropical forest on the Guiana Shield, a globally important forest carbon reservoir. We model recovery of timber and carbon with a breakpoint regression that allowed us to capture elevated tree mortality immediately after logging. Recovery rates of timber and carbon were governed by the presence of residual trees (i.e., trees that persisted through the first harvest). The liberation treatment stimulated faster recovery of timber albeit at a carbon cost. Model results suggest a threshold logging intensity beyond which forests managed for timber and carbon derive few benefits from RIL, with recruitment and residual growth not sufficient to offset losses. Inclusion of the breakpoint at which carbon and timber gains outpaced postlogging mortality led to high predictive accuracy, including out‐of‐sample R² values >90%, and enabled inference on demographic changes postlogging. Our modeling framework is broadly applicable to studies that aim to quantify impacts of logging on forest recovery. Overall, we demonstrate that initial mortality drives variation in recovery rates, that the second harvest depends on old growth wood, and that timber intensification lowers carbon stocks.     

Assessing the effects of selective logging on birds in Neotropical piedmont and cloud montane forests

In tropical and subtropical forests there is limited information about how to integrate sustainable timber management with the conservation of biodiversity. We examined the effect of selective logging on the bird community to help develop management guidelines to assure the conservation of biodiversity in forests managed for timber production. The study design consisted of control and harvested plots in piedmont and cloud forests of the subtropical montane forests of the Andes in northwestern Argentina. We conducted bird point-count surveys combined with distance estimation. Breeding season bird community composition was more similar between control and logged forest in both the cloud forest and piedmont, than between the two elevations, probably because Neotropical bird communities change dramatically along elevational gradients. Within each elevation zone, community composition changed significantly between harvested and control forests. Both between and within each elevation zone no significant differences in bird density were detected. Similarly, when we analyzed bird density according to diet guilds no general pattern could be extracted. However, we found a significantly greater density of cavity nesters and lower of non-cavity nesters in control plots, probably because most trees that can develop suitable cavities were extracted in logged plots and these plots had a greater structural diversity enabling more nesting resources. Grouping species according to their nesting habitat requirements has rarely been used in the neotropics and other tropical and subtropical forests, but focusing management attention on cavity nesters might address the most sensitive portion of the avian community as well as other species dependent on trees likely to hold cavities.     

Multiple stages of tree seedling recruitment are altered in tropical forests degraded by selective logging

Tropical forest degradation is a global environmental issue. In degraded forests, seedling recruitment of canopy trees is vital for forest regeneration and recovery. We investigated how selective logging, a pervasive driver of tropical forest degradation, impacts canopy tree seedling recruitment, focusing on an endemic dipterocarp Dryobalanops lanceolata in Sabah, Borneo. During a mast‐fruiting event in intensively logged and nearby unlogged forest, we examined four stages of the seedling recruitment process: seed production, seed predation, and negative density‐dependent germination and seedling survival. Our results suggest that each stage of the seedling recruitment process is altered in logged forest. The seed crop of D. lanceolata trees in logged forest was one‐third smaller than that produced by trees in unlogged forest. The functional role of vertebrates in seed predation increased in logged forest while that of non‐vertebrates declined. Seeds in logged forest were less likely to germinate than those in unlogged forest. Germination increased with local‐scale conspecific seed density in unlogged forest, but seedling survival tended to decline. However, both germination and seedling survival increased with local‐scale conspecific seed density in logged forest. Notably, seed crop size, germination, and seedling survival tended to increase for larger trees in both unlogged and logged forests, suggesting that sustainable timber extraction and silvicultural practices designed to minimize damage to the residual stand are important to prevent seedling recruitment failure. Overall, these impacts sustained by several aspects of seedling recruitment in a mast‐fruiting year suggest that intensive selective logging may affect long‐term population dynamics of D. lanceolata. It is necessary to establish if other dipterocarp species, many of which are threatened by the timber trade, are similarly affected in tropical forests degraded by intensive selective logging.     

Diversity and community composition of ectomycorrhizal fungi in a dry deciduous dipterocarp forest in Thailand

Large forest areas of South-East Asia, are dominated by the Dipterocarpaceae tree family, which contains many important timber species. Unlike many other tropical trees, Dipterocarpaceae rely on ectomycorrhizal (ECM) root symbiosis for their mineral nutrition. This study aims to document the richness and community composition of ECM fungi in a dry deciduous forest in Thailand. Combining morphological and molecular identification methods revealed 69 species of ECM fungi that belong to 17 phylogenetic lineages. The /russula-lactarius, /tomentella-thelephora, /sordariales, /sebacina and /cantharellus lineages were the most species-rich. The fungal richness is comparable to other tropical rain forest sites, but the phylogenetic community structure has elements of both tropical and temperate ecosystems. Unlike tropical rain forests, the Cenococcum geophilum complex was one of the most frequent fungal taxa that had a relatively high ITS genetic diversity over the small sampling area. This study provides the first snapshot insight into the fungal community of dry dipterocarp forests. However, it is necessary to broaden the spatial and temporal scales of sampling to improve our understanding of the below-ground relations of dry and humid tropical forests.     

Impacts of Selective Logging and Agricultural Clearing on Forest Structure, Floristic Composition and Diversity, and Timber Tree Regeneration in the Ituri Forest, Democratic Republic of Congo

Mature tropical forests at agricultural frontiers are of global conservation concern as the leading edge of global deforestation. In the Ituri Forest of DRC, as in other tropical forest areas, road creation associated with selective logging results in spontaneous human colonization, leading to the clearing of mature forest for agricultural purposes. Following 1-3 years of cultivation, farmlands are left fallow for periods that may exceed 20 years, resulting in extensive secondary forest areas impacted by both selective logging and swidden agriculture. In this study, we assessed forest structure, tree species composition and diversity and the regeneration of timber trees in secondary forest stands (5-10 and ~40 years old), selectively logged forest stands, and undisturbed forests at two sites in the Ituri region. Stem density was lower in old secondary forests (~40 years old) than in either young secondary or mature forests. Overall tree diversity did not significantly differ between forest types, but the diversity of trees ≥10 cm dbh was substantially lower in young secondary forest stands than in old secondary or mature forests. The species composition of secondary forests differed from that of mature forests, with the dominant Caesalpinoid legume species of mature forests poorly represented in secondary forests. However, in spite of prior logging, the regeneration of high value timber trees such as African mahoganies (Khaya anthotheca and Entandrophragma spp.) was at least 10 times greater in young secondary forests than in mature forests. We argue that, if properly managed and protected, secondary forests, even those impacted by both selective logging and small-scale shifting agriculture, may have high potential conservation and economic value.     

Documenting Biogeographical Patterns of African Timber Species Using Herbarium Records: A Conservation Perspective Based on Native Trees from Angola

In many tropical regions the development of informed conservation strategies is hindered by a dearth of biodiversity information. Biological collections can help to overcome this problem, by providing baseline information to guide research and conservation efforts. This study focuses on the timber trees of Angola, combining herbarium (2670 records) and bibliographic data to identify the main timber species, document biogeographic patterns and identify conservation priorities. The study recognized 18 key species, most of which are threatened or near-threatened globally, or lack formal conservation assessments. Biogeographical analysis reveals three groups of species associated with the enclave of Cabinda and northwest Angola, which occur primarily in Guineo-Congolian rainforests, and evergreen forests and woodlands. The fourth group is widespread across the country, and is mostly associated with dry forests. There is little correspondence between the spatial pattern of species groups and the ecoregions adopted by WWF, suggesting that these may not provide an adequate basis for conservation planning for Angolan timber trees. Eight of the species evaluated should be given high conservation priority since they are of global conservation concern, they have very restricted distributions in Angola, their historical collection localities are largely outside protected areas and they may be under increasing logging pressure. High conservation priority was also attributed to another three species that have a large proportion of their global range concentrated in Angola and that occur in dry forests where deforestation rates are high. Our results suggest that timber tree species in Angola may be under increasing risk, thus calling for efforts to promote their conservation and sustainable exploitation. The study also highlights the importance of studying historic herbarium collections in poorly explored regions of the tropics, though new field surveys remain a priority to update historical information.     

Selective logging in tropical forests decreases the robustness of liana–tree interaction networks to the loss of host tree species

Selective logging is one of the major drivers of tropical forest degradation, causing important shifts in species composition. Whether such changes modify interactions between species and the networks in which they are embedded remain fundamental questions to assess the ‘health’ and ecosystem functionality of logged forests. We focus on interactions between lianas and their tree hosts within primary and selectively logged forests in the biodiversity hotspot of Malaysian Borneo. We found that lianas were more abundant, had higher species richness, and different species compositions in logged than in primary forests. Logged forests showed heavier liana loads disparately affecting slow-growing tree species, which could exacerbate the loss of timber value and carbon storage already associated with logging. Moreover, simulation scenarios of host tree local species loss indicated that logging might decrease the robustness of liana–tree interaction networks if heavily infested trees (i.e. the most connected ones) were more likely to disappear. This effect is partially mitigated in the short term by the colonization of host trees by a greater diversity of liana species within logged forests, yet this might not compensate for the loss of preferred tree hosts in the long term. As a consequence, species interaction networks may show a lagged response to disturbance, which may trigger sudden collapses in species richness and ecosystem function in response to additional disturbances, representing a new type of ‘extinction debt’.     

Western lowland gorilla density and nesting behavior in a Gabonese forest logged for 25 years: implications for gorilla conservation

The tropical forests of the Congo Basin constitute biodiversity refuges that still hold large numbers of species, including endemic and endangered vertebrates. Along with several key species, the critically endangered western lowland gorilla (WLG) potentially contributes to forest dynamics through seed dispersal. Considering the extensive influence of timber harvesting on tropical forest ecosystems, the survival of gorilla populations in logged forests might prove critical for forest ecosystem conservation. We estimated WLG density, through a nest count survey, in a forest in southeast Gabon that has been logged for 25 years. Nesting behavior and habitat use were described and we applied generalized linear models to identify the factors that influence gorilla day and night habitat use. The estimated density of weaned gorillas, 1.5 gorillas km^?2, is comparable with estimates from some protected areas and other sustainably managed sites within their range. Habitat type had the greatest influence on nest site distribution. We observed a preference for nesting in open terra firma forest, and open habitats in general, which supports the findings of previous studies. Habitat use during the day was strongly influenced by habitat type and human activities, and to a lesser degree by functional and non-functional roads, and rivers. Our results support the suggestion that logged forests are suitable habitats for WLG if hunting and poaching are controlled. We recommend collaborations between timber operators and scientists to improve the conservation potential of tropical forests and enhance the wildlife-management aspects of logging practices.     

Phylogenetic Age is Positively Correlated with Sensitivity to Timber Harvest in Bornean Mammals

The reasons that forest vertebrates differ in their response to selective timber extraction in tropical forests remain poorly characterized. Understanding what determines response and sensitivity can indicate how forest management might yield greater conservation benefits, and help us identify which lesser-known species may be especially vulnerable. We assessed the response of 41 Bornean mammals to selective timber harvest and tested eight hypotheses regarding the correlation between those responses and a range of species characteristics. Multivariate analyses show that phylogenetic species age is a key variable determining sensitivity. Older species are less able to cope with the effects of selective timber harvest. Most of these species are endemic to insular southeast Asia, and do not occur on the Asian mainland. These species are more specialized, and appear less able to cope with habitat change. In contrast, species tolerant to logging evolved more recently. This group tends to be omnivorous or herbivorous, to use all vegetation strata, and to be regionally widespread. This finding allows the sensitivity to habitat disturbance of lesser-known species to be predicted, and therefore has important conservation implications. These new insights also help in the design of large-scale forest landscapes that combine sustainable forest management and species conservation requirements. We recognize that these functions can be compatible, but that some species still need completely protected areas for their survival.     

Changes of community composition at multiple trophic levels due to hunting in Nigerian tropical forests

Hunting in tropical forests decimates large mammals, and this may have direct and indirect effects on other trophic levels and lead to trophic cascades. We compared replicated sites of hunted and protected forests in southeastern Nigeria, with respect to community composition of primates, other mammals, birds, plant seedlings, and mature trees. We make predictions regarding the community composition at the different trophic levels. In forests where large primates are rare, we hypothesize that their ecological role will not be fully compensated for by small frugivores. We apply multivariate methods to assess changes in community composition of mammals, birds, and seedlings, controlling for any differences between sites in the other groups, including mature trees. Medium and large (4–180 kg) primates were much rarer in hunted sites, while porcupine and rock hyrax increased in abundance with hunting. In contrast, the community composition of birds was similar in both types of forests. Seedling communities were significantly related to the community composition of mammals, and thus strongly affected by hunting. In protected forests primate dispersed plant seedling species dominated, whereas in hunted forests the seedling community was shifted towards one dominated by abiotically dispersed species. This was probably both a consequence of reduced seed dispersal by primates, and increased seed predation by rodents and hyrax. Hence we found no evidence for buffering effects on tree regeneration through functional compensation by non‐hunted animals (such as birds). Our results highlight how seedling communities are changed by the complex plant–animal intera ctions, triggered by the loss of seed dispersers. The results predict a rarity of primate‐dispersed trees in future tropical forest canopies; a forest less diverse in timber and non‐timber resources.     

Predicting which tropical tree species are vulnerable to forest disturbances

Tropical forest management often focuses on a few high‐value timber species because they are thought to be the most vulnerable in logged forests. However, other tree species may be vulnerable to secondary effects of logging, like loss of vertebrate dispersers. We examined vulnerability of tree species to loss of vertebrate dispersers in Mabira, a heavily disturbed tropical rainforest in Uganda. Fruit characteristics and shade tolerance regimes of 269 tree species were compiled. Stem densities of tree species producing fruits of various sizes and having different shade tolerance regimes were computed for Mabira and compared with densities of conspecifics in Budongo, a less disturbed forest with similar floral composition. Seventy per cent of tree species in Mabira are animal‐dispersed, of which 10% are large‐fruited light demanders. These species are the most vulnerable because they rarely recruit beneath adult conspecifics and are exclusively dispersed by large vertebrates, also vulnerable in heavily disturbed forests. Comparison of densities between Mabira and Budongo showed that large‐fruited light demanders had a lower density in Mabira. Other categories of tree species had similar densities in both forests. It is plausible that the low density of large‐fruited light demanders is due to limited recruitment caused by dispersal limitations.     

The Tropical managed Forests Observatory: a research network addressing the future of tropical logged forests

While attention on logging in the tropics has been increasing, studies on the long-term effects of silviculture on forest dynamics and ecology remain scare and spatially limited. Indeed, most of our knowledge on tropical forests arises from studies carried out in undisturbed tropical forests. This bias is problematic given that logged and disturbed tropical forests are now covering a larger area than the so-called primary forests. A new network of permanent sample plots in logged forests, the Tropical managed Forests Observatory (TmFO), aims to fill this gap by providing unprecedented opportunities to examine long-term data on the resilience of logged tropical forests at regional and global scales. TmFO currently includes 24 experimental sites distributed across three tropical regions, with a total of 490 permanent plots and 921 ha of forest inventories.     

Long-Term Impacts of Fuelwood Extraction on a Tropical Montane Cloud Forest

Fuelwood extracted from natural forests serves as a principal energy source in rural regions of many tropical countries. Although fuelwood extraction (even low intensities) might strongly impact the structure and species composition of natural forests, long-term studies remain scarce. Here, we estimate the potential long-term impacts (over several hundred years) of such repeated harvesting of single trees on tropical montane cloud forest in central Veracruz, Mexico, by applying a process-based forest growth model. We simulate a wide range of possible harvesting scenarios differing in wood volume harvested and preferred tree species and sizes, and use a set of indicators to compare their impacts on forest size structure and community composition. Results showed that the overall impact on forest structure and community composition increased linearly with the amount of harvested wood volume. Even at low levels of harvesting, forest size structure became more homogeneous in the long term because large old trees disappeared from the forest, but these changes might take decades or even centuries. Although recruitment of harvested species benefited from harvesting, species composition shifted to tree species that are not used for fuelwood. Our results demonstrate that fuelwood extraction can have marked long-term impacts on tropical montane cloud forests. The results also offer the possibility to support the design of management strategies for the natural species-rich forests that achieve a balance between economic needs and ecological goals of the stakeholders. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.