Avian Habitat Preference in Tropical Forest Restoration in Southern Costa Rica

An important question for tropical forest restoration is whether degraded lands can be actively managed to attract birds. We censused birds and measured vegetation structure at 27 stations in young (6–9‐yr old) actively and passively restored pasture and old growth forest at Las Cruces Biological Station in southern Costa Rica. During 481 10‐min point counts, we detected a high diversity—186 species—of birds using the restoration area. Surprisingly, species richness and detection frequency did not differ among habitats, and proportional similarity of bird assemblages to old growth forest did not differ between restoration treatments. Bird detection frequency was instead explained by exotic grass cover and understory stem density—vegetation structures that were not strongly impacted by active restoration. The similarity of bird assemblages in actively and passively restored forest may be attributed to differential habitat preferences within and among feeding guilds, low structural contrast between treatments, or the effect of nucleation from actively restored plots into passively restored areas. Rapid recovery of vegetation in this recently restored site is likely due to its proximity to old growth forest and the lack of barriers to effective seed dispersal. Previous restoration studies in highly binary environments (i.e., open pasture vs. tree plantation) have found strong differences in bird abundance and richness. Our data contradict this trend, and suggest that tropical restoration ecologists should carefully consider: (1) when the benefits of active restoration outweigh the cost of implementation; and (2) which avian guilds should be used to measure restoration success given differential responses to habitat structure.     

The use of spatially patterned methods for vegetation restoration and management across systems

Widespread degradation of natural lands has created an urgent need for restoration. However, the high cost of conventional techniques limits the extent and success of restoration efforts. As a result, practitioners have developed new cost‐effective techniques. Spatially patterned restoration methods, where established clusters of plant species serve as propagule sources across a broad target area, have been proposed as practical restoration techniques. The spatial patterning is expected to reduce initial costs and provide ecological benefits such as increasing habitat heterogeneity. Over the past three decades, multiple spatially patterned restoration methods have emerged around the globe; however, it is unclear whether applications and theoretical foundations have been connected across methods. We conducted a literature review and bibliometric network analyses to (1) examine patterns in focal study systems, cost‐effectiveness, and ecological outcomes for spatially patterned restoration methods and (2) analyze connectivity among the bodies of literature associated with common spatially patterned restoration methods to identify knowledge gaps and synergies. We found the three most commonly studied methods are applied nucleation, slot seeding, and strip seeding. Applied nucleation studies mainly occurred in tropical forests and emphasized plant diversity and seed‐dispersing animal visitation. Slot‐seeding and strip‐seeding studies both primarily occurred in temperate grasslands and emphasized plant establishment and production. Applied nucleation and slot‐seeding approaches had distinct theoretical bases, as evidenced by patterns in reference citation, while strip‐seeding approaches did not draw from a unified body of literature. We discuss the need for full economic analyses and theoretical links between the different methods.     

Fire effects and ecological recovery pathways of tropical montane cloud forests along a time chronosequence

Tropical montane cloud forests (TMCFs) harbour high levels of biodiversity and large carbon stocks. Their location at high elevations make them especially sensitive to climate change, because a warming climate is enhancing upslope species migration, but human disturbance (especially fire) may in many cases be pushing the treeline downslope. TMCFs are increasingly being affected by fire, and the long‐term effects of fire are still unknown. Here, we present a 28‐year chronosequence to assess the effects of fire and recovery pathways of burned TMCFs, with a detailed analysis of carbon stocks, forest structure and diversity. We assessed rates of change of carbon (C) stock pools, forest structure and tree‐size distribution pathways and tested several hypotheses regarding metabolic scaling theory (MST), C recovery and biodiversity. We found four different C stock recovery pathways depending on the selected C pool and time since last fire, with a recovery of total C stocks but not of aboveground C stocks. In terms of forest structure, there was an increase in the number of small stems in the burned forests up to 5–9 years after fire because of regeneration patterns, but no differences on larger trees between burned and unburned plots in the long term. In support of MST, after fire, forest structure appears to approximate steady‐state size distribution in less than 30 years. However, our results also provide new evidence that the species recovery of TMCF after fire is idiosyncratic and follows multiple pathways. While fire increased species richness, it also enhanced species dissimilarity with geographical distance. This is the first study to report a long‐term chronosequence of recovery pathways to fire suggesting faster recovery rates than previously reported, but at the expense of biodiversity and aboveground C stocks.     

Recovery of mammal diversity in tropical forests: a functional approach to measuring restoration

Ecological restoration is increasingly applied in tropical forests to mitigate biodiversity loss and recover ecosystem functions. In restoration ecology, functional richness, rather than species richness, often determines community assembly, and measures of functional diversity provide a mechanistic link between diversity and ecological functioning of restored habitat. Vertebrate animals are important for ecosystem functioning. Here, we examine the functional diversity of small‐to‐medium sized mammals to evaluate the diversity and functional recovery of tropical rainforest. We assess how mammal species diversity and composition and functional diversity and composition, vary along a restoration chronosequence from degraded pasture to “old‐growth” tropical rainforest in the Wet Tropics of Australia. Species richness, diversity, evenness, and abundance did not vary, but total mammal biomass and mean species body mass increased with restoration age. Species composition in restoration forests converged on the composition of old‐growth rainforest and diverged from pasture with increasing restoration age. Functional metrics provided a clearer pattern of recovery than traditional species metrics, with most functional metrics significantly increasing with restoration age when taxonomic‐based metrics did not. Functional evenness and dispersion increased significantly with restoration age, suggesting that niche complementarity enhances species' abundances in restored sites. The change in community composition represented a functional shift from invasive, herbivorous, terrestrial habitat generalists and open environment specialists in pasture and young restoration sites, to predominantly endemic, folivorous, arboreal, and fossorial forest species in older restoration sites. This shift has positive implications for conservation and demonstrates the potential of tropical forest restoration to recover rainforest‐like, diverse faunal communities.     

Resilience and restoration of tropical and subtropical grasslands,savannas, and grassy woodlands

Despite growing recognition of the conservation values of grassy biomes, our understanding of how to maintain and restore biodiverse tropical grasslands (including savannas and open‐canopy grassy woodlands) remains limited. To incorporate grasslands into large‐scale restoration efforts, we synthesised existing ecological knowledge of tropical grassland resilience and approaches to plant community restoration. Tropical grassland plant communities are resilient to, and often dependent on, the endogenous disturbances with which they evolved – frequent fires and native megafaunal herbivory. In stark contrast, tropical grasslands are extremely vulnerable to human‐caused exogenous disturbances, particularly those that alter soils and destroy belowground biomass (e.g. tillage agriculture, surface mining); tropical grassland restoration after severe soil disturbances is expensive and rarely achieves management targets. Where grasslands have been degraded by altered disturbance regimes (e.g. fire exclusion), exotic plant invasions, or afforestation, restoration efforts can recreate vegetation structure (i.e. historical tree density and herbaceous ground cover), but species‐diverse plant communities, including endemic species, are slow to recover. Complicating plant‐community restoration efforts, many tropical grassland species, particularly those that invest in underground storage organs, are difficult to propagate and re‐establish. To guide restoration decisions, we draw on the old‐growth grassland concept, the novel ecosystem concept, and theory regarding tree cover along resource gradients in savannas to propose a conceptual framework that classifies tropical grasslands into three broad ecosystem states. These states are: (1) old‐growth grasslands (i.e. ancient, biodiverse grassy ecosystems), where management should focus on the maintenance of disturbance regimes; (2) hybrid grasslands, where restoration should emphasise a return towards the old‐growth state; and (3) novel ecosystems, where the magnitude of environmental change (i.e. a shift to an alternative ecosystem state) or the socioecological context preclude a return to historical conditions.     

Large‐sized rare tree species contribute disproportionately to functional diversity in resource acquisition in African tropical forest

Increasing evidence is available for a positive effect of biodiversity on ecosystem productivity and standing biomass, also in highly diverse systems as tropical forests. Biodiversity conservation could therefore be a critical aspect of climate mitigation policies. There is, however, limited understanding of the role of individual species for this relationship, which could aid in focusing conservation efforts and forest management planning. This study characterizes the functional specialization and redundancy for 95% of all tree species (basal area weighted percentage) in a diverse tropical forest in the central Congo Basin and relates this to species' abundance, contribution to aboveground carbon, and maximum size. Functional characterization is based on a set of traits related to resource acquisition (wood density, specific leaf area, leaf carbon, nitrogen and phosphorus content, and leaf stable carbon isotope composition). We show that within both mixed and monodominant tropical forest ecosystems, the highest functional specialization and lowest functional redundancy are solely found in rare tree species and significantly more in rare species holding large‐sized individuals. Rare species cover the entire range of low and high functional redundancy, contributing both unique and redundant functions. Loss of species supporting functional redundancy could be buffered by other species in the community, including more abundant species. This is not the case for species supporting high functional specialization and low functional redundancy, which would need specific conservation attention. In terms of tropical forest management planning, we argue that specific conservation of large‐sized trees is imperative for long‐term maintenance of ecosystem functioning.     

Is applied nucleation a straightforward,cost‐effective forest restoration approach? Counter‐response to Holl and Zahawi (2018)

Holl and Zahawi (2018) agree on a number of approaches that we describe in a local case study of tropical montane cloud forest restoration in Mexico. However, they contend our argument that most applied nucleation projects have taken place in tropical lowlands as a mistake. They also provide data on the per‐hectare cost of restoration projects in Costa Rica and compare it to a higher cost figure of plantation‐style reforestation, a view contrary to ours. Last, Holl and Zahawi recognize that applied nucleation requires specialized personnel, but imply that the amount of training needed for this endeavor is not very different than what is required to implement other forest recovery projects. In this counter‐response, we provide some refinements to our original opinion and offer further information in support to our perspective.     

Demographic costs and benefits of natural regeneration during tropical forest restoration

For tropical forest restoration to result in long‐term biodiversity gains, native trees must establish self‐sustaining populations in degraded sites. While many have asked how seedling recruitment varies between restoration treatments, the long‐term fate of these recruits remains unknown. We address this research gap by tracking natural recruits of 27 species during the first 7 years of a tropical forest restoration experiment that included both planted and naturally regenerating plots. We used an individual‐based model to estimate the probability that a seedling achieves reproductive maturity after several years of growth and survival. We found an advantage for recruits in naturally regenerating plots, with up to 40% increased probability of reproduction in this treatment, relative to planted plots. The demographic advantage of natural regeneration was highest for mid‐successional species, with relatively minor differences between treatments for early‐successional species. Our research demonstrates the consequences of restoration decision making across the life cycle of tropical tree species.     

Applied nucleation is a straightforward,cost‐effective forest restoration approach: reply to Ramírez‐Soto et al. (2018)

We agree with Ramírez‐Soto and colleagues that applied nucleation can be an effective approach for tropical forest restoration both in lowland and higher elevation tropical forests. We also contend that it is cheaper than standard plantation‐style plantings and is straightforward to train personnel to implement this approach.     

Putting it back: Woody debris in young restoration plantings to stimulate return of reptiles

Despite active investment in restoration, some habitat features can be slow to develop on formerly degraded land and can consequently pose persistent barriers to the re‐establishment of specialist species. Coarse woody debris (CWD) is a critical resource for a whole suite of animal taxa but remains an underappreciated component of some forest ecosystems and restoration activities. The extent to which recovery of animal communities can be accelerated through artificial supplementation of woody debris is poorly understood especially for highly diverse tropical forest systems. Here, we report early results from an experiment designed to manipulate CWD in young restoration plantings (0–7 year old) in tropical north‐east Australia for the purposes of facilitating re‐establishment of rainforest reptiles. After 1 year, we demonstrate that CWD addition within restoration plantings adjacent to remnant forest can increase the local abundance of reptiles and promote colonisation of the log‐specialist Prickly Skink (Gnypetoscincus queenslandiae). These preliminary results, however, are based on observations of just 44 individual reptiles encompassing seven species. Ongoing monitoring will elucidate longer‐term outcomes to enable a proper evaluation of when and where CWD addition might be most beneficial in realising restoration goals.     

Improving Planting Stocks for the Brazilian Atlantic Forest Restoration through Community‐Based Seed Harvesting Strategies

High‐diversity reforestation can help jumpstart tropical forest restoration, but obtaining viable seedlings is a major constraint: if nurseries do not offer them, it is hard to plant all the species one would like. From 2007 to 2009, we investigated five different seed acquisition strategies employed by a well‐established tree nursery in southeastern Brazil, namely (1) in‐house seed harvesters; (2) hiring a professional harvester; (3) amateur seed harvesters; or (4) a seed production cooperative, as well as (5) participating in a seed exchange program. In addition, we evaluated two strategies not dependent on seeds: harvesting seedlings from native tree species found regenerating under Eucalyptus plantations, and in a native forest remnant. A total of 344 native tree and shrub species were collected as seeds or seedlings, including 2,465 seed lots. Among these, a subset of 120 species was obtained through seed harvesting in each year. Overall, combining several strategies for obtaining planting stocks was an effective way to increase species richness, representation of some functional groups (dispersal syndromes, planting group, and shade tolerance), and genetic diversity of seedlings produced in forest tree nurseries. Such outcomes are greatly desirable to support high‐diversity reforestation as part of tropical forest restoration. In addition, community‐based seed harvesting strategies fostered greater socioeconomic integration of traditional communities in restoration projects and programs, which is an important bottleneck for the advance of ecological restoration, especially in developing countries. Finally, we discuss some of the limitations of the various strategies for obtaining planting stocks and the way forward for their improvement.     

Canopy Size Dependent Facilitations from the Native Shrub Rhodomyrtus tomentosa to the Early Establishment of Native Trees Castanopsis fissa and Syzygium hancei in Tropical China

The quantitative role of the canopy size of nurse shrubs on microenvironment and native tree establishment in degraded tropical lands has been seldom studied. In a 21‐month field experiment, we aimed to test the effect of a native shrub with different canopy sizes on the early establishment of native trees as part of the effort of forest restoration in tropical China. We examined the microenvironment, and the seedling establishment and growth of two native trees: Castanopsis fissa and Syzygium hancei in both open space (OS) microsite and microsite under the canopy of the native pioneer shrub Rhodomyrtus tomentosa. Shrub microsite was further divided into large canopy (LC), and medium canopy (MC) microsite, based on the shrub leaf area indices. Results showed that relative to OS, LC had higher soil nutrient concentration and water content, and lower photosynthetic active radiation (PAR), while MC had lower PAR and higher soil exchangeable Mg, K, and Ca. Survival and growth were mostly enhanced, while water stress and photoinhibition reduced for C. fissa seedlings in MC and S. hancei seedlings in LC. It is found that the beneficial effects of the native shrub on seedling establishment and growth result mostly from the improvement in nutrient and water availabilities, the reduction in plant stress caused by harsh summer light, and the specific ecological requirements of different tree species. We suggest that different canopy sizes of native shrub R. tomentosa may be explored to target different native trees and hence promote forest restoration in degraded tropical ecosystems.     

A Strategy for Restoration of Montane Forest in Anthropogenic Fern Thickets in the Dominican Republic

Deforested tropical areas are often colonized by competitive ferns that inhibit forest succession. In thickets of such a fern (Dicranopteris pectinata), we investigated methods for initiating restoration of tropical montane forest in the Ébano Verde Scientific Reserve (Dominican Republic). In clearings in the thickets, growth and survivorship of 18 common early‐ and late‐successional woody species were tested, with and without fertilizer (poultry litter). Three years after sowing, life history did not affect survivorship, but early‐successional species grew faster than late‐successional species (height increase 153 ± 103 cm vs. 81 ± 67 cm [mean ± 1 SD]). Inga fagifolia, a late‐successional species, and Alchornea latifolia, an early‐successional species, had 160 ± 62 cm mean height increase, and low mortality rates (<4%). In contrast, four late‐successional species (Cyrilla racemiflora, Myrcia deflexa, Prestoea acuminata var. montana, and Mora abbottii), and one early‐successional species, Ocotea leucoxylon, had approximately 39% mortality and height increase of 43 ± 48 cm. Brunellia comocladifolia had high mortality (55%), but averaged 340 ± 201 cm height increase, and was the only species whose growth was improved by fertilization. Fertilization improved survivorship of only one species, Piper aduncum. After three years, soils in the clearings had low pH and available P and did not differ significantly from soils in thickets. However, based on the growth rates and survivorship of sown woody plants, these soils did not appear to be a barrier for restoration. Although a complementary study demonstrated substantial natural regeneration, active planting should be used to increase plant density and diversity, especially where natural regeneration is poor.     

Community‐based native seed production for restoration in Brazil – the role of science and policy

• Large‐scale restoration programmes in the tropics require large volumes of high quality, genetically diverse and locally adapted seeds from a large number of species. However, scarcity of native seeds is a critical restriction to achieve restoration targets.
• In this paper, we analyse three successful community‐based networks that supply native seeds and seedlings for Brazilian Amazon and Cerrado restoration projects. In addition, we propose directions to promote local participation, legal, technical and commercialisation issues for up‐scaling the market of native seeds for restoration with high quality and social justice.
• We argue that effective community‐based restoration arrangements should follow some principles: (i) seed production must be based on real market demand; (ii) non‐governmental and governmental organisations have a key role in supporting local organisation, legal requirements and selling processes; (iii) local ecological knowledge and labour should be valued, enabling local communities to promote large‐scale seed production; (iv) applied research can help develop appropriate techniques and solve technical issues.
• The case studies from Brazil and principles presented here can be useful for the up‐scaling restoration ecology efforts in many other parts of the world and especially in tropical countries where improving rural community income is a strategy for biodiversity conservation and restoration.

     

No evidence for consistent long‐term growth stimulation of 13 tropical tree species: results from tree‐ring analysis

The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate–vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO₂‐fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest‐stand level, but insights into species‐specific growth changes – that ultimately determine community‐level responses – are lacking. Here, we analyse species‐specific growth changes on a centennial scale, using growth data from tree‐ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size‐class isolation) growth‐trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8–10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large‐scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO₂ would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.     

The hydroclimatic and ecophysiological basis of cloud forest distributions under current and projected climates

Background

Tropical montane cloud forests (TMCFs) are characterized by a unique set of biological and hydroclimatic features, including frequent and/or persistent fog, cool temperatures, and high biodiversity and endemism. These forests are one of the most vulnerable ecosystems to climate change given their small geographic range, high endemism and dependence on a rare microclimatic envelope. The frequency of atmospheric water deficits for some TMCFs is likely to increase in the future, but the consequences for the integrity and distribution of these ecosystems are uncertain. In order to investigate plant and ecosystem responses to climate change, we need to know how TMCF species function in response to current climate, which factors shape function and ecology most and how these will change into the future.

Scope

This review focuses on recent advances in ecophysiological research of TMCF plants to establish a link between TMCF hydrometeorological conditions and vegetation distribution, functioning and survival. The hydraulic characteristics of TMCF trees are discussed, together with the prevalence and ecological consequences of foliar uptake of fog water (FWU) in TMCFs, a key process that allows efficient acquisition of water during cloud immersion periods, minimizing water deficits and favouring survival of species prone to drought-induced hydraulic failure.

Conclusions

Fog occurrence is the single most important microclimatic feature affecting the distribution and function of TMCF plants. Plants in TMCFs are very vulnerable to drought (possessing a small hydraulic safety margin), and the presence of fog and FWU minimizes the occurrence of tree water deficits and thus favours the survival of TMCF trees where such deficits may occur. Characterizing the interplay between microclimatic dynamics and plant water relations is key to foster more realistic projections about climate change effects on TMCF functioning and distribution.     

Effects of Tree Guards and Weed Mats on the Establishment of Native Tree Seedlings: Implications for Forest Restoration in Hong Kong, China

The effects of tree guards and weed mats on establishment and growth of native tree seedlings, Thick‐leaved oak (Cyclobalanopsis edithiae (Skan) Schott., Fagaceae), planted in an exposed hillside grassland in Hong Kong, were investigated. The natural regeneration of C. edithiae is poor due to a lack of seed dispersal agents and high seed predation, and therefore, this species is often targeted for forest restoration. The experiment lasted for 3.5 years during which the height, basal diameter, and crown diameter of individual seedlings were measured and survivorship recorded. The use of weed mats alone did not have a significant effect, but a combination of tree guards and weed mats led to a significant improvement in establishment, survivorship, and growth of the seedlings during the experimental period. Initially, the guards promoted rapid height growth of the seedlings, although lateral growth and secondary stem thickening were compromised. After the seedlings grew over the tree guards, the basal diameter and crown diameter increased at a notably faster rate. The combined effect of the tree guard and weed mat on the seedling growth pattern was found to be beneficial and contributed to the high survivorship of the seedlings. Comparing the survivorship data and the costs of various treatments, the use of tree guards in combination with the weed mats was demonstrated to be more cost‐effective than planting the seedlings without tree guards or weed mats. The potential for applying the technique in afforestation programs with native tree species for forest restoration in Hong Kong and other tropical regions is discussed.     

Jump‐starting coastal wetland restoration: a comparison of marsh and mangrove foundation species

During coastal wetland restoration, foundation plant species are critical in creating habitat, modulating ecosystem functions, and supporting ecological communities. Following initial hydrologic restoration, foundation plant species can help stabilize sediments and jump‐start ecosystem development. Different foundation species, however, have different traits and environmental tolerances. To understand how these traits and tolerances impact restoration trajectories, there is a need for comparative studies among foundation species. In subtropical and tropical climates, coastal wetland restoration practitioners can sometimes choose between salt marsh and/or mangrove foundation species. Here, we compared the early life history traits and environmental tolerances of two foundation species: (1) a salt marsh grass (Spartina alterniflora) and (2) a mangrove tree (Avicennia germinans). In an 18‐month study of a recently restored coastal wetland in southeastern Louisiana (USA), we examined growth and survival along an elevation gradient and compared expansion and recruitment rates. We found that the rapid growth, expansion, and recruitment rates of the salt marsh grass make it a better species for quickly establishing ecological structure at suitable elevations. The slower growth, limited expansion, and lower recruitment of the mangrove species show its restricted capacity for immediate structural restoration, especially in areas where it co‐occurs with perennial salt marsh species. Our findings suggest that the structural attributes needed in recently restored areas can be achieved sooner using fast‐growing foundation species. Following salt marsh grass establishment, mangroves can then be used to further assist ecosystem development. This work highlights how appropriate foundation species can help jump‐start ecosystem development to meet restoration objectives.     

Low‐cost agricultural waste accelerates tropical forest regeneration

Lower‐cost tropical forest restoration methods, particularly those framed as win–win business‐protected area partnerships, could dramatically increase the scale of tropical forest restoration activities, thereby providing a variety of societal and ecosystem benefits, including slowing both global biodiversity loss and climate change. Here we describe the long‐term regenerative effects of a direct application of agricultural waste on tropical dry forest. In 1998, as part of an innovative agricultural waste disposal service contract, an estimated 12,000 Mg of processed orange peels and pulp were applied to a 3 ha portion of a former cattle pasture with compacted, rocky, nutrient‐poor soils characteristic of prolonged fire‐based land management and overgrazing in Área de Conservación Guanacaste, northwestern Costa Rica. After 16 years, the experimental plot showed a threefold increase in woody plant species richness, a tripling of tree species evenness (Shannon Index), and a 176% increase in aboveground woody biomass over an adjacent control plot. Hemispheric photography showed significant increases in canopy closure in the area where orange waste was applied relative to control. Orange waste deposition significantly elevated levels of soil macronutrients and important micronutrients in samples taken 2 and 16 years after initial orange waste application. Our results point to promising opportunities for valuable synergisms between agricultural waste disposal and tropical forest restoration and carbon sequestration.     

Light Response of Seedlings of a Central African Timber Tree Species,Lophira alata (Ochnaceae), and the Definition of Light Requirements

Light is of primary importance in structuring tropical tree communities. Light exposure at seedling and adult stages has been used to characterize the ecological profile of tropical trees, with many implications in forest management and restoration ecology. Most shade‐tolerance classification systems have been proposed based on empirical observations in a specific area and thus result in contradictions among categories assigned to a given species. In this study, we aimed to quantify the light requirements for seedling growth of a Central African timber tree, Lophira alata (Ochnaceae), taking into account effects of population origin. In two controlled experiments: a light response experiment and a comparative population experiment, conducted in southwestern Cameroon, using seeds collected from four populations (three from Cameroon and one from Gabon), we examined the quantitative responses to irradiance of seedlings. After 2 years, mortality was very low (<3%), even in extremely low irradiance. Growth and biomass allocation patterns varied in response to light, with intermediate irradiance (24–43%) providing optimal conditions. Light response differed between populations. The Boumba population in the northeastern edge of the species' distribution exhibited the highest light requirements, suggesting a local adaptation. As a result of positive growth at low irradiance and maximum growth at intermediate irradiance, we concluded that L. alata exhibits characteristics of both non‐pioneer and pioneer species. Implications of our results to propose an objective way to assign the light requirement for tropical tree species are discussed.