Colonization of woody seedlings in the understory of actively and passively restored tropical moist forests

The status of woody seedling colonization gives clues about the self‐sustainability of restored forests, a tenet of restoration success. Little is known about woody seedling colonization in restored afrotropical forests. We evaluated effects of restoration methods (active vs. passive), sampling year, restoration age, and distance from old‐growth forests on seedling colonization in restored afrotropical moist forests. Seedlings were measured in 2011 and 2014 in 71 clusters of 284 permanent sampling plots (12.6 m² each) in actively (initially 3–16 years old) and 21 clusters of 63 plots in passively restored forests (initially 16 years old) in Kibale National Park, western Uganda. Seedlings were also measured in nearby old‐growth forests in three clusters of five plots in 2014. We determined species diversity, richness, abundance per plot, and species composition as measures of seedling colonization in restored and old‐growth forests. We found that diversity, richness, and abundance of seedlings were significantly higher in passively than actively restored forests. Diversity and richness but not abundance significantly increased between sampling years and with restoration age. Distance from old‐growth forests did not significantly affect diversity, richness, and abundance. Species composition of actively and passively restored forests was different from that of old‐growth forests after 19 years since restoration started. Our results show that passive restoration should be the preferred method for recovering afrotropical forests, and highlight the effect of continued management on biodiversity of restored forests.     

Impacts of wind disturbance on fragmented tropical forests: A review and synthesis

Abstract Wind disturbance is an important ecological force in the tropics, especially in the cyclonic and hurricane zones from about 7–20° latitude. Damage from intense winds may be especially severe in fragmented forests because of their abrupt artificial margins and denuded surrounding landscapes. We review available information on the effects of windstorms on fragmented forests, synthesizing studies from Australasia, Amazonia and elsewhere in the tropics. Wind damage in fragmented landscapes can be influenced by a range of factors, such as forest‐edge orientation, edge structure, the size of nearby clearings and local topography. We argue that wind disturbances are likely to interact with, and exacerbate, a range of deleterious environmental changes in fragmented forests. Among the most important of these are altered forest structure, shifts in plant species composition, exotic‐plant invasions, reduced carbon storage and elevated vulnerability to fire. The damaging impacts of winds on fragmented forests could potentially increase in the future, particularly if global warming leads to increasingly severe or frequent windstorms.     

Managing subsistence hunting in the changing landscape of Neotropical rain forests

Subsistence hunting has been a vital activity for local people across Neotropical rain forests (NRF). While providing a reliable source of protein, subsistence hunting also reflected the strong relationships that connected local people with the species and ecosystems in which they inhabited. However, the social and ecological context in which subsistence hunting can be sustainable has been altered. The relatively small groups that hunted in large and mostly undisturbed forests, using traditional weapons, have been replaced by a growing population using fragmented habitats and modern hunting methods. Thus, subsistence hunting is less likely to be sustainable, threatening the food security of local people and the persistence of species with critical roles in the functioning of NRF. Managing subsistence hunting in this changing context will require a more efficient combination of tools which might include banning the hunting of large and sensitive species, strengthening protected areas, alternatives to reduce the role of wildlife protein on local people's subsistence and, in some cases, voluntary resettlements of local people, from areas that could still be used as refuge for endangered species.     

Impacts of anthropogenic climate change on tropical montane forests: an appraisal of the evidence

In spite of their small global area and restricted distributions, tropical montane forests (TMFs) are biodiversity hotspots and important ecosystem services providers, but are also highly vulnerable to climate change. To protect and preserve these ecosystems better, it is crucial to inform the design and implementation of conservation policies with the best available scientific evidence, and to identify knowledge gaps and future research needs. We conducted a systematic review and an appraisal of evidence quality to assess the impacts of climate change on TMFs. We identified several skews and shortcomings. Experimental study designs with controls and long-term (≥10 years) data sets provide the most reliable evidence, but were rare and gave an incomplete understanding of climate change impacts on TMFs. Most studies were based on predictive modelling approaches, short-term (<10 years) and cross-sectional study designs. Although these methods provide moderate to circumstantial evidence, they can advance our understanding on climate change effects. Current evidence suggests that increasing temperatures and rising cloud levels have caused distributional shifts (mainly upslope) of montane biota, leading to alterations in biodiversity and ecological functions. Neotropical TMFs were the best studied, thus the knowledge derived there can serve as a proxy for climate change responses in under-studied regions elsewhere. Most studies focused on vascular plants, birds, amphibians and insects, with other taxonomic groups poorly represented. Most ecological studies were conducted at species or community levels, with a marked paucity of genetic studies, limiting understanding of the adaptive capacity of TMF biota. We thus highlight the long-term need to widen the methodological, thematic and geographical scope of studies on TMFs under climate change to address these uncertainties. In the short term, however, in-depth research in well-studied regions and advances in computer modelling approaches offer the most reliable sources of information for expeditious conservation action for these threatened forests.     

Comparing seed removal rates in actively and passively restored tropical moist forests

High rates of seed removal can impede forest recovery, but tropical seed removal studies are few and mainly from the neotropics. Little is known about the comparative influences of active restoration (i.e. planting) and passive restoration (i.e. protection of natural regrowth) on seed removal. We conducted an evaluation of seed removal in grasslands, natural forests (tropical moist semideciduous forest), and actively (21‐, 17‐, 16‐, 11‐, 8‐, and 6‐year‐old) and passively (21‐year‐old) restored forests in Kibale National Park, Uganda. We wanted to compare the effect of vegetation type, time since restoration and restoration actions (i.e. active vs. passive) on removal of seeds of five animal‐dispersed tree species during wet and dry seasons. Seeds were either fully exposed or placed in closed mesh cages or under a mesh roof. We used differential removal rates between these treatments to attribute seed removal to different animal taxa. Seed removal rate (percentage of seed removed over a 4‐day period) was highest in passively restored forests, compared with actively restored forests, grasslands, and natural forests. We detected no significant relationship between time since restoration and seed removal rates within actively restored sites. Seed removal rate from roofed treatments was not significantly different from removal from open treatments but was significantly higher than removal from closed treatments, which we interpret as reflecting the greater effect of small mammals versus insects. Smaller seeds tended to be removed at a greater rate than larger seeds. We discuss the implications of these findings for forest regeneration.     

Drought effects on seedling survival in a tropical moist forest

The amount and seasonality of rainfall varies strongly in the tropics, and plant species abundance, distribution and diversity are correlated with rainfall. Drought periods leading to plant stress occur not only in dry forests, but also in moist and even wet forests. We quantified experimentally the effect of drought on survival of first year seedlings of 28 co-occurring tropical woody plant species in the understory of a tropical moist forest. The seedlings were transplanted to plots and subjected to a drought and an irrigation treatment for 22 weeks during the dry season. Drought effects on mortality and wilting behavior varied greatly among species, so that relative survival in the dry treatment ranged from 0% to about 100% of that in the irrigated treatment. Drought stress was the main factor in mortality, causing about 90% (median) of the total mortality observed in the dry treatment. In almost half of the species, the difference in survival between treatments was not significant even after 22 weeks, implying that many of the species are well adapted to drought in this forest. Relative drought survival was significantly higher in species associated with dry habitats than in those associated with wet habitats, and in species with higher abundance on the dry side of the Isthmus of Panama, than in those more abundant on the wet side. These data show that differential species survival in response to drought, combined with variation in soil moisture availability, may be important for species distribution at the local and regional scale in many tropical forests.     

Genetic diversity of ectomycorrhizal Basidiomycetes from African and Indian tropical rain forests

Ectomycorrhizal (ECM) fungi have a worldwide distribution. However, the ecology of tropical ECM fungi is poorly documented, limiting our understanding of the symbiotic associations between tropical plants and fungi. ECM Basidiomycete diversity was investigated for the first time in two tropical rain forests in Africa (Western Upper Guinea) and in Asia (Western Ghats, India), using a fragment of the mitochondrial large subunit rRNA gene to type 140 sporocarps and 54 ectomycorrhizas. To evaluate taxonomic diversity, phylogenetic analyses were performed, and 40 sequences included from identified European specimens were used as taxonomic benchmarks. Five clades were recovered corresponding to six taxonomic groups: boletoids, sclerodermatoids, russuloids, thelephoroids, and a clade grouping the Amanitaceae and Tricholomataceae families. Our results revealed that the Russulaceae species display a great diversity with several putative new species, especially in Guinea. Other taxonomic issues at family/section levels are also briefly discussed. This study provides preliminary insights into taxonomic diversity, ECM status, and biogeographic patterns of ECM fungi in tropical two rain forest ecosystems, which appear to be as diverse as in temperate and boreal forests.     

Importance of natural licks for the mammals in Bornean inland tropical rain forests

An intensive camera-trapping study and a nutrient analysis were carried out to understand how natural licks are important for mammals in inland tropical rain forests where soil cations are usually depleted. Using camera traps, we investigated the fauna, food habits, and the frequency of visitation by species at five natural licks in the Deramakot forest reserve, Sabah, Malaysia. All food-habit types of mammals (carnivore, herbivore-frugivore, insectivore, and omnivore), which included 29 (78.4%) of 37 species known in Deramakot, were recorded at the natural licks. The sambar deer, followed by the bearded pig, the lesser mouse-deer, the Malay badger, and the orangutan were the most commonly recorded species and represented 77.5% in terms of the frequency of appearance in all photographs taken throughout the year. These results indicated that, although the proportion of species recorded at the natural licks relative to the whole mammalian fauna of the forest was high, the frequency of visitation greatly varied among the species, and only a few species dominated. The frequency of visitation seemed to reflect both the density of species and the demand for the minerals, because some endangered, low-density species were more frequently recorded by cameras than expected—for example, the orangutan which was one of the top five species among natural-lick users. The natural licks with greater concentrations of minerals in seepage soil water were significantly preferred by the sambar deer and the bearded pig than those with lower concentrations of minerals. This result suggests that the chemical properties of soil water in natural licks determine the frequency of visitation of these herbivorous species that have strong demand for minerals.     

A comprehensive synthesis of liana removal experiments in tropical forests

Lianas are a quintessential feature of tropical forests and are often perceived as being poorly studied. However, liana removal studies may be one of the most common experimental manipulations in tropical forest ecology. In this review, we synthesize data from 64 tropical liana removal experiments conducted over the past 90 yr. We explore the direction and magnitude of the effects of lianas on tree establishment, growth, survival, reproduction, biomass accretion, and plant and animal diversity in ecological and forestry studies. We discuss the geographical biases of liana removal studies and compare the various methods used to manipulate lianas. Overall, we found that lianas have a clear negative effect on trees, and trees benefitted from removing lianas in nearly every study across all forest types. Liana cutting significantly increased light and water availability, and trees responded with vastly greater reproduction, growth, survival, and biomass accumulation compared to controls where lianas were present. Removing lianas during logging significantly reduced damage of future merchantable trees and improved timber production. Our review demonstrates that lianas have an unequivocally detrimental effect on every metric of tree performance measured, regardless of forest type, forest age, or geographic location. However, lianas also appear to have a positive contribution to overall forest plant diversity and to different animal groups. Therefore, managing lianas reduces logging damage and improves timber production; however, the removal lianas may also have a negative effect on the faunal community, which could ultimately harm the plant community.     

Estimating aboveground net biomass change for tropical and subtropical forests: Refinement of IPCC default rates using forest plot data

As countries advance in greenhouse gas (GHG) accounting for climate change mitigation, consistent estimates of aboveground net biomass change (?AGB) are needed. Countries with limited forest monitoring capabilities in the tropics and subtropics rely on IPCC 2006 default ?AGB rates, which are values per ecological zone, per continent. Similarly, research into forest biomass change at a large scale also makes use of these rates. IPCC 2006 default rates come from a handful of studies, provide no uncertainty indications and do not distinguish between older secondary forests and old‐growth forests. As part of the 2019 Refinement to the 2006 IPCC Guidelines for National Greenhouse Gas Inventories, we incorporate ?AGB data available from 2006 onwards, comprising 176 chronosequences in secondary forests and 536 permanent plots in old‐growth and managed/logged forests located in 42 countries in Africa, North and South America and Asia. We generated ?AGB rate estimates for younger secondary forests (≤20 years), older secondary forests (>20 years and up to 100 years) and old‐growth forests, and accounted for uncertainties in our estimates. In tropical rainforests, for which data availability was the highest, our ?AGB rate estimates ranged from 3.4 (Asia) to 7.6 (Africa) Mg ha^?1 year^?1 in younger secondary forests, from 2.3 (North and South America) to 3.5 (Africa) Mg ha^?1 year^?1 in older secondary forests, and 0.7 (Asia) to 1.3 (Africa) Mg ha^?1 year^?1 in old‐growth forests. We provide a rigorous and traceable refinement of the IPCC 2006 default rates in tropical and subtropical ecological zones, and identify which areas require more research on ?AGB. In this respect, this study should be considered as an important step towards quantifying the role of tropical and subtropical forests as carbon sinks with higher accuracy; our new rates can be used for large‐scale GHG accounting by governmental bodies, nongovernmental organizations and in scientific research.     

Bryophytes in perennially moist forests of Papua New Guinea: ecological orientation and predictions of disturbance effects

Water relations of bryophytes must be understood along at least four dimensions: hydration/dehydration frequency; hydration duration; dehydration duration; degree of water loss. All these are biomass-dependent functions such that large colonies of bryophytes may maintain hydration longer than smaller colonies. The spread of a bryophyte colony allows lateral movement of capillary water; contiguous clones will thus allow lateral conduction of that water. In contrast, separated tufts or cushions may store water but will contribute very little to its transfer over the surface of the phorophyte.
Many of the tropical rainforests of New Guinea have a larger mass of epiphytic bryophytes than any temperate forest. The primary effect of disturbance in these forests is a reduction in bryomass, presumably due to desiccation because of increased insolation and wind movement. The reduction of bryomass will decrease water and mineral retention in the bryophyte clones; and the reduction in the size of those clones will reduce their contiguity and thus interfere with bryophyte-mediatcd water and mineral transfer. Disturbance may alter bryophyte species frequency but, under present agricultural and forestry practices in New Guinea, I have seen no evidence of significant loss of bryophyte species.     

The FORMNET‐B database: monitoring the biomass and dynamics of disturbed and degraded tropical forests

The biomass and dynamics of disturbed and degraded tropical forests have mostly been ignored in the recent scientific literature, partly because of a spotlight on old‐growth forests but also due to a lack of long‐term data from degraded forests. There is a pressing need to understand the rates and patterns of growth, mortality and recruitment in degraded forests, not only because they are increasing in area relative to old‐growth forests, but also due to their potential capacity to sequester large amounts of carbon dioxide from the atmosphere. This paper introduces a permanent forest plot network and database, FORMNET‐B (GIVD ID# NA‐BZ‐001), designed to study the long‐term dynamics of disturbed and degraded tropical forests in Belize, Central America.     

Impacts of climate variability on tree demography in second growth tropical forests: the importance of regional context for predicting successional trajectories

Naturally regenerating and restored second growth forests account for over 70% of tropical forest cover and provide key ecosystem services. Understanding climate change impacts on successional trajectories of these ecosystems is critical for developing effective large‐scale forest landscape restoration (FLR) programs. Differences in environmental conditions, species composition, dynamics, and landscape context from old growth forests may exacerbate climate impacts on second growth stands. We compile data from 112 studies on the effects of natural climate variability, including warming, droughts, fires, and cyclonic storms, on demography and dynamics of second growth forest trees and identify variation in forest responses across biomes, regions, and landscapes. Across studies, drought decreases tree growth, survival, and recruitment, particularly during early succession, but the effects of temperature remain unexplored. Shifts in the frequency and severity of disturbance alter successional trajectories and increase the extent of second growth forests. Vulnerability to climate extremes is generally inversely related to long‐term exposure, which varies with historical climate and biogeography. The majority of studies, however, have been conducted in the Neotropics hindering generalization. Effects of fire and cyclonic storms often lead to positive feedbacks, increasing vulnerability to climate extremes and subsequent disturbance. Fragmentation increases forests’ vulnerability to fires, wind, and drought, while land use and other human activities influence the frequency and intensity of fire, potentially retarding succession. Comparative studies of climate effects on tropical forest succession across biogeographic regions are required to forecast the response of tropical forest landscapes to future climates and to implement effective FLR policies and programs in these landscapes.     

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.     

Sampling bees in tropical forests and agroecosystems: a review

Bees are the predominant pollinating taxa, providing a critical ecosystem service upon which many angiosperms rely for successful reproduction. Available data suggests that bee populations worldwide are declining, but scarce data in tropical regions precludes assessing their status and distribution, impact on ecological services, and response to management actions. Herein, we reviewed >150 papers that used six common sampling methods (pan traps, baits, Malaise traps, sweep nets, timed observations and aspirators) to better understand their strengths and weaknesses, and help guide method selection to meet research objectives and development of multi-species monitoring approaches. Several studies evaluated the effectiveness of sweep nets, pan traps, and malaise traps, but only one evaluated timed observations, and none evaluated aspirators. Only five studies compared two or more of the remaining four sampling methods to each other. There was little consensus regarding which method would be most reliable for sampling multiple species. However, we recommend that if the objective of the study is to estimate abundance or species richness, malaise traps, pan traps and sweep nets are the most effective sampling protocols in open tropical systems; conversely, malaise traps, nets and baits may be the most effective in forests. Declining bee populations emphasize the critical need in method standardization and reporting precision. Moreover, we recommend reporting a catchability coefficient, a measure of the interaction between the resource (bee) abundance and catching effort. Melittologists could also consider existing methods, such as occupancy models, to quantify changes in distribution and abundance after modeling heterogeneity in trapping probability, and consider the possibility of developing monitoring frameworks that draw from multiple sources of data.     

Fragment shape and tree species composition in tropical forests: a landscape level investigation

Fragmentation of tropical forest alters community composition as a result of changes in forest shape. This paper uses 22 hypotheses to test the effect of fragment shape on tree species composition in Ghana, West Africa, within biological categories of regeneration guild, rarity, phenology and dispersal. For both regenerating and mature trees, relationships between species composition and the shape of forest fragments were complex; almost half were significant but many failed to support the established hypotheses. Irregular shaped fragments had high proportions of regenerating, light‐demanding pioneers and mature, animal‐dispersed species. Species common to Ghana formed the foundation of communities in fragments of all shapes. Investigation at the landscape level indicated broad patterns of species change. Rigorous hypothesis testing is needed, following extensive demographic work on the ground, before population dynamics within tropical forest fragments can be comprehended fully and applied to conservation management.     

The myriad consequences of hunting for vertebrates and plants in tropical forests

Humans hunt forest vertebrates throughout the tropics. Many preferred game species consume flowers, fruit, seeds and/or leaves, and these interactions will cause their harvest to ramify through forests. Three related issues will determine how severely the harvest of forest vertebrates influences the plant community. First, the types of species selected by hunters and the intensity of the harvest will determine which vertebrates are removed and which remain. Second, the possible presence of ecologically similar, non-game species able to expand their activities to fulfill the ecological role of heavily exploited species will determine how severely the harvest disrupts ecological relationships between the community of forest vertebrates and the community of forest plants. Finally, hunters will alter plant species composition if the harvest of vertebrates differentially affects mutualists or pests of particular plant species. Hunters will also alter plant diversity if the harvest of vertebrates disrupts ecological mechanisms that permit plant species to coexist. I examine hunter selectivity, the intensity of the hunt, possible compensation by non-game species, and the types and strengths of interactions among game species and plants for tropical forests to determine when and where these outcomes occur.