Canopy phylogenetic, chemical and spectral assembly in a lowland Amazonian forest

? Canopy chemistry and spectroscopy offer insight into community assembly and ecosystem processes in high-diversity tropical forests, but phylogenetic and environmental factors controlling chemical traits underpinning spectral signatures remain poorly understood. ? We measured 21 leaf chemical traits and spectroscopic signatures of 594 canopy individuals on high-fertility Inceptisols and low-fertility Ultisols in a lowland Amazonian forest. The spectranomics approach, which explicitly connects phylogenetic, chemical and spectral patterns in tropical canopies, provided the basis for analysis. ? Intracrown and intraspecific variation in chemical traits varied from 1.4 to 36.7% (median 9.3%), depending upon the chemical constituent. Principal components analysis showed that 14 orthogonal combinations were required to explain 95% of the variation among 21 traits, indicating the high dimensionality of canopy chemical signatures among taxa. Inceptisols and lianas were associated with high leaf nutrient concentrations and low concentrations of defense compounds. Independent of soils or plant habit, an average 70% (maximum 89%) of chemical trait variation was explained by taxonomy. At least 10 traits were quantitatively linked to remotely sensed signatures, which provided highly accurate species classification. ? The results suggest that taxa found on fertile soils carry chemical portfolios with a deep evolutionary history, whereas taxa found on low-fertility soils have undergone trait evolution at the species level. Spectranomics provides a new connection between remote sensing and community assembly theory in high-diversity tropical canopies.     

海南尖峰岭热带山地雨林林冠层树种功能多样性特征

以海南尖峰岭热带山地雨林3块1 hm²样地为研究对象,利用11个林冠功能性状结合样地地形及林冠乔木树种样地清查数据,分别基于单维性状和多维性状比较物种多度加权对群落功能离散度指数——平均成对距离(MPD)和平均最近类群距离(MNTD)的影响;同时分析林冠层功能丰富度(FRic)与物种丰富度之间的关系,最后利用零模型探讨不同生境类型下标准化效应MPD和MNTD(经过物种多度加权且剔除群落物种丰富度差异影响)的变化,进而评价林冠层群落水平功能多样性格局及其对局域生境异质性的响应.结果表明: 功能性状维度和物种多度对MPD的影响强烈,不同维度功能性状多度加权前后MPD相关性较弱(R=0.359～0.628);但对MNTD的影响相对较弱,不同维度功能性状多度加权前后MNTD相关性较强(R=0.746～0.820);未经物种多度加权的MPD和MNTD均普遍高估了林冠层的功能离散度.林冠层功能丰富度与物种丰富度基本呈指数相关关系(F=128.20;R²=0.632;AIC=97.72;P＜0.001),且功能丰富度很有可能存在一定的物种丰富度阈值.基于不同维度功能性状的林冠层功能多样性格局及其生境响应存在一定的差异性.在生物竞争激烈的低沟生境中,林冠层功能多样性倾向于比预期零模型随机产生的功能多样性高,林冠树种功能性状表现出离散分布;而在其他生境类型中,林冠层功能多样性倾向于接近或低于随机产生的功能多样性,林冠树种功能性状随机或聚集分布.     

Detecting invisible growth rings of trees in seasonally dry forests in Thailand: isotopic and wood anatomical approaches

We measured radial variation of carbon isotope composition and vessel traits in tree species in seasonally dry forests of Northeast Thailand to explore a more reliable and amenable method of tropical dendrochronology for trees that lack visually detectable and consistent growth rings. Six Dipterocarpaceae species (3 Shorea, 2 Dipterocarpus, and 1 Hopea species) with indistinct or irregular growth rings and teak (Tectona grandis), a species which forms distinct growth rings, were examined. The δ¹³C value variations in all species showed annual cyclicity. Dipterocarpaceae species usually marked the lowest values of δ¹³C in the middle of the growing season, whereas teak had the lowest values at nearly the end of the growing season. Since the growing season of the species examined almost corresponds to the rainy season in the study area, the δ¹³C variation was likely caused by the change in moisture availability. The different variation pattern of teak was attributable to its stronger dependence on ¹³C-enriched reserved material early in the growing season. Changes in tree vessel traits for all species examined also showed annual cyclicity. Dipterocarpaceae species showed significant correlation between δ¹³C values and vessel measurements. Vessel lumen (mean area, tangential and radial diameter, and proportion of total area) had a negative correlation, whereas vessel frequency showed a positive correlation. The correlations indicated that changes in vessel traits were caused by the seasonal variation of moisture available to the trees. Thus, we concluded that methods using wood anatomy, as well as δ¹³C, have great potential for use as tools in tropical dendrochronology within the context of seasonal climate.     

Convergent elevation trends in canopy chemical traits of tropical forests

The functional biogeography of tropical forests is expressed in foliar chemicals that are key physiologically based predictors of plant adaptation to changing environmental conditions including climate. However, understanding the degree to which environmental filters sort the canopy chemical characteristics of forest canopies remains a challenge. Here, we report on the elevation and soil‐type dependence of forest canopy chemistry among 75 compositionally and environmentally distinct forests in nine regions, with a total of 7819 individual trees representing 3246 species collected, identified and assayed for foliar traits. We assessed whether there are consistent relationships between canopy chemical traits and both elevation and soil type, and evaluated the general role of phylogeny in mediating patterns of canopy traits within and across communities. Chemical trait variation and partitioning suggested a general model based on four interconnected findings. First, geographic variation at the soil‐Order level, expressing broad changes in fertility, underpins major shifts in foliar phosphorus (P) and calcium (Ca). Second, elevation‐dependent shifts in average community leaf dry mass per area (LMA), chlorophyll, and carbon allocation (including nonstructural carbohydrates) are most strongly correlated with changes in foliar Ca. Third, chemical diversity within communities is driven by differences between species rather than by plasticity within species. Finally, elevation‐ and soil‐dependent changes in N, LMA and leaf carbon allocation are mediated by canopy compositional turnover, whereas foliar P and Ca are driven more by changes in site conditions than by phylogeny. Our findings have broad implications for understanding the global ecology of humid tropical forests, and their functional responses to changing climate.     

Genetic variation in nine Shorea species (Dipterocarpaceae) in Indonesia revealed by AFLPs

Shorea is the largest and most important genus of the Dipterocarpaceae. The genetic diversity and structure of nine Shorea species from two different locations, namely Nanjak Makmur in Sumatra and Sumalindo in Borneo, were evaluated using amplified fragment length polymorphism (AFLP) markers. A total of 274 trees were investigated at 85 polymorphic AFLP loci. Levels of genetic diversity of these species ranged from  = 0.100 for S. acuminata to  = 0.165 for S. blumutensis. The population of rare species S. blumutensis possessed the highest genetic diversity suggesting that geographically restricted species can have levels of genetic variation comparable to closely related widespread common congeners. Analyses of molecular variance revealed that the genetic variation was mainly found among species in both locations (57.7% in Sumatra; 56.3% in Borneo). The unweighted pairgroup method using arithmetic averages dendrogram of all samples revealed an almost complete separation of species. Thus, AFLP markers proved appropriate for phylogenetic studies of Shorea species. Specific markers have been detected showing high-frequency differences among species and between regions within species. Sequence information of these markers can be used to develop specific polymerase chain reaction markers for wood identification. The possibility of interspecific hybridization was discussed.     

Structure and dynamics in seasonal dry evergreen forest in northeastern Thailand

Abstract. Studies of tropical forest dynamics have often been based on one large-scale permanent plot, representative of a given forest type. Broad classifications of tropical forest types are expected to include a wide range of stand structures, dynamics patterns and species compositions – a range which cannot be represented in a single plot. To demonstrate this problem two 1-ha permanent plots, dominated by Hopea ferrea and Shorea henryana (both Dipterocarpaceae), respectively, were established in 1987 in seasonal dry evergreen forest at the Sakaerat Environmental Research Station in northeastern Thailand. In 1997 the plots were remeasured as to patterns of recruitment, mortality and growth. The Hopea plot was relatively static with low mortality, recruitment and growth. The Shorea plot was very dynamic with high rates of growth, mortality and recruitment. If the current trends continue, the plots are likely to further diverge. Even if the study of a large forest plot provides a good insight into tropical forest dynamics, it is necessary to consider the entire local pattern of variation.     

Effects of <Emphasis Type="Italic">Morella faya</Emphasis> tree invasion on aboveground carbon storage in Hawaii

The negative effects of biological invasion are often the focus of ecological studies, but few have considered potential positive impacts, such as increased carbon storage, resulting from invasion. We combined airborne imaging spectrometer and LiDAR (light detection and ranging) observations with field measurements to assess if the highly invasive nitrogen-fixing tree Morella faya alters canopy 3-D structure and aboveground biomass (AGB) along a 1,500 mm precipitation gradient in Hawaii. Airborne analysis of canopy water content, leaf nitrogen concentration, fractional canopy cover, and vegetation height facilitated mapping of native- and Morella-dominated canopies in rainforest, woodland–savanna and shrubland ecosystems, with Morella detection errors ranging from 0 to 13.4%. Allometric equations were developed to relate the combined LiDAR and spectral data to field-based AGB estimates (r ² = 0.97, P < 0.01), and to produce a map of biomass stocks throughout native and invaded ecosystems. The structure of the invasive Morella canopies varied by ecosystem type, and the invader shaded out native understory plants in rainforest zones. Despite a 350% increase in AGB going from shrubland to rainforest, Morella did not increase average AGB in any ecosystem it invaded. Furthermore, spatial distributions of biomass indicated that Morella decreased maximum AGB in the woodland–savanna ecosystems. We conclude that Morella tree invasion does not enhance aboveground carbon stocks in any ecosystem it invades in Hawaii, thereby minimizing its contribution to this potentially important ecosystem service. We also found that the fusion of spectral and LiDAR remote sensing provided canopy chemical and structural data facilitating a landscape assessment of how biological invasion alters on carbon stocks and other ecosystem properties.     

Borneo Illipe,a fat product from differentShorea spp. (Dipterocarpaceae)

Fat extracted from kernels of Shorea spp. (Dipterocarpaceae) in the tropical rainforests of Borneo and Sumatra are traded under the name Borneo Illipe. The fat is used in chocolate formulations. The majority of kernels collected are fromShorea species in the sectionPachycarpae. Fat characteristics ofShorea species in this section show differences in fat quality in relation to its use as exotic sources in Cocoa Butter Equivalent (CBE) formulations. In order to comment on the future potential of the naturally occurring tree populations distribution maps of fiveShorea spp. involved in Borneo Illipe utilization are presented.     

Relationships among Amphibian Assemblage Structure,Wetland pH,and Forest Cover

A common mesofilter approach to conservation of biological diversity and ecosystem function used in agricultural and urban landscapes is maintenance of wetlands and an undisturbed terrestrial buffer surrounding wetlands. Although it is generally accepted that forest buffers protect wetland-associated biological diversity and ecosystem function, the effectiveness and optimal spatial extent of buffers is still an area of debate. During 2007 and 2008 we surveyed amphibians and environmental conditions associated with 54 depression wetlands on the Delmarva Peninsula of Maryland, USA, to examine the role of forest buffers and wetland characteristics in structuring amphibian communities. Forest cover within a 50-m buffer surrounding wetlands was correlated (r = −0.81) with wetland pH but no other wetland characteristics. Wetland pH, canopy cover, hydroperiod, and adjacent forest cover were important predictors of wetland use by individual amphibian species, with many species more likely to occur at wetlands that dried late in the hydrological year and with open canopies. At least one common species preferred circumneutral pH and several restricted-distribution species preferred lower pH (<5). Contrary to expectations, relationships between species occurrence and adjacent forest cover were negative. Our results suggest that current regulations that provide buffers of 30 m or less do not provide adequate protection of wetland water chemistry but that forest encroachment into wetlands may be a threat to the integrity of amphibian communities and should be the target of monitoring, future research, and management efforts. © 2021 The Wildlife Society.     

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.     

Phylogenetic Analysis of Dipterocarps Using Random Amplified Polymorphic DNA Markers

The phylogenetic relationships among 12 species belonging tothree different genera (Shorea, HopeaandAnisoptera) of Dipterocarpaceaewere studied using random amplified polymorphic DNA (RAPD) markers.A modified CTAB DNA extraction protocol was used to obtain tannin-and polysaccharide-free genomic DNA from mature leaves. Clusteranalysis of data from six random primers placed the 12 speciesin three groups corresponding to their respective genera. Fourdistinct nodes ofShoreaspp. and two ofHopeaspp. could be identified.Anisopteramegistocarpaserved as an outgroup, and was unique when comparedto the other genera examined. RAPD profiles of five individualsofH. odoratawith six random primers were identical, suggestingthat there is little intraspecific variation in this species.The RAPD technique can thus be successfully applied for thestudy of phylogenetic relationships of this important groupof tropical timber trees.Copyright 1998 Annals of Botany Company Dipterocarpaceae,Anisopteraspp.,Hopeaspp.,Shoreaspp., RAPD markers, tropical timber trees.     

Defining a spectrum of integrative trait‐based vegetation canopy structural types

Vegetation canopy structure is a fundamental characteristic of terrestrial ecosystems that defines vegetation types and drives ecosystem functioning. We use the multivariate structural trait composition of vegetation canopies to classify ecosystems within a global canopy structure spectrum. Across the temperate forest sub‐set of this spectrum, we assess gradients in canopy structural traits, characterise canopy structural types (CST) and evaluate drivers and functional consequences of canopy structural variation. We derive CSTs from multivariate canopy structure data, illustrating variation along three primary structural axes and resolution into six largely distinct and functionally relevant CSTs. Our results illustrate that within‐ecosystem successional processes and disturbance legacies can produce variation in canopy structure similar to that associated with sub‐continental variation in forest types and eco‐climatic zones. The potential to classify ecosystems into CSTs based on suites of structural traits represents an important advance in understanding and modelling structure–function relationships in vegetated ecosystems.     

Molecular phylogeny of Dipterocarpaceae in Southeast Asia using RFLP of PCR-amplified chloroplast genes

Dipterocarpaceae is the dominant family of Southeast Asia's climax tropical rain forest region, and it contains the region's most important commercial timber species. A molecular phylogeny of the Dipterocarpaceae subfamily Dipterocapoideae was constructed using restriction fragment length polymorphisms of polymerase chain reaction-amplified specific genes in chloroplast DNA. A total of 141 site changes were detected among ten genera and 30 species in 11 different genes: rbcL, psbA, psbD, rpoB, rpoC, petB, atpH, 16S, psaA, petA and trnK. Phylogenetic trees constructed by Wanger parsimony and neighbor-joining methods, using Upuna as the outgroup, displayed five monophytelic groups that included Upuna: HopeaShorea-Parashorea-Neobalanocarpus; Dryobalanops; Dipterocarpus; Anisoptera-Vatica-Cotylelobium; and Upuna. The phylogenetic trees clearly separate species with two different base chromosome numbers: the first group is x=7, and the other is x=11. The x=7 group is thought to be in a synapomorphic character state. Parashorea lucida is a sister to most Shorea species. Neobalanocarpus heimii and Hopea from a clade of a sister to two Shorea species, and Cotylelobium and Vatica are closely related species. Our conclusions agree with a phylogeny derived from wood anatomy data analysis, and with Symington's and Ashton's taxonomic classifications.The raw data of the PCR-RFLP analysis can be obtained from the authors     

Influence of Seasonal Variations in Soil Water Availability on Gas Exchange of Tropical Canopy Trees

Seasonal variations in environmental conditions influence the functioning of the whole ecosystem of tropical rain forests, but as yet little is known about how such variations directly influence the leaf gas exchange and transpiration of individual canopy tree species. We examined the influence of seasonal variations in relative extractable water in the upper soil layers on predawn leaf water potential, saturated net photosynthesis, leaf dark respiration, stomatal conductance, and tree transpiration of 13 tropical rain forest canopy trees (eight species) over 2 yr in French Guiana. The canopies were accessed by climbing ropes attached to the trees and to a tower. Our results indicate that a small proportion of the studied trees were unaffected by soil water depletion during seasonal dry periods, probably thanks to efficient deep root systems. The trees showing decreased tree water status (i.e., predawn leaf water potential) displayed a wide range of leaf gas exchange responses. Some trees strongly regulated photosynthesis and transpiration when relative extractable water decreased drastically. In contrast, other trees showed little variation, thus indicating good adaptation to soil drought conditions. These results have important applications to modeling approaches: indeed, precise evaluation and grouping of these response patterns are required before any tree‐based functional models can efficiently describe the response of tropical rain forest ecosystems to future changes in environmental conditions.     

Horizontal, but not vertical canopy structure is related to stand functional diversity in a subtropical slope forest

The aim of this study was to analyse the relation of horizontal and vertical canopy structure to tree functional diversity of a highly diverse subtropical broad-leaved slope forest, stratified for different successional stages. This is of particular interest because many key ecosystem processes and functions are related to the arrangement of forest canopies. We assessed the effect of stand-related functional diversity (FD_Q, measured as Rao’s quadratic entropy of leaf traits), together with other environmental variables on horizontal [measured as relative crown projection areas (CPA_r)] and vertical [relative crown overlap, coefficients of variation (CV) of crown positioning variables] structure of the upper canopy at the local neighbourhood level. The analyses with mixed effects models revealed a negative relation (p = 0.025; estimate −0.07) between FD_Q and CPA_r. No significant effect of FD_Q on vertical canopy structure has been found (p > 0.05). The findings are discussed with regard to resource partitioning and niche differentiation of canopy and sub-canopy species. Successional stage positively impacted the CV of crown length (p = 0.019; estimate 0.03) but did not affect other response variables. The sloping terrain strongly influenced vertical canopy structure as revealed by the significant effect of slope inclination on CV of crown length (p = 0.004; estimate −0.05) and of slope aspect on CV of mean crown height (p = 0.036; estimate −0.03). The high complexity of vertical crown positioning depending on the heterogeneous sloping terrain of the study area may have obscured relations of FD_Q to vertical canopy structure.     

Diverse Optical and Photosynthetic Properties in a Neotropical Dry Forest during the Dry Season: Implications for Remote Estimation of Photosynthesis1

Using optical and photosynthetic assays from a canopy access crane, we examined the photosynthetic performance of tropical dry forest canopies during the dry season in Parque Metropolitano, Panama City, Panama. Photosynthetic gas exchange, chlorophyll fluorescence, and three indices derived from spectral reflectance (the normalized difference vegetation index, the simple ratio, and the photochemical reflectance index) were used as indicators of structural and physiological components of photosynthetic activity. Considerable interspecific variation was evident in structural and physiological behavior in this forest stand, which included varying degrees of foliage loss, altered leaf orientation, stomatal closure, and photosystem II downregulation. The normalized difference vegetation index and the simple ratio were closely related to canopy structure and absorbed radiation for most species, but failed to capture the widely divergent photosynthetic behavior among evergreen species exhibiting various degrees of downregulation. The photochemical reflectance index and chlorophyll fluorescence were related indicators of photosynthetic downregulation, which was not detectable with the normalized difference vegetation index or simple ratio. These results suggest that remote sensing methods that ignore downregulation cannot capture within‐stand variability in actual carbon flux for this diverse forest type. Instead, these findings support a sampling approach that derives photosynthetic fluxes from a consideration of both canopy light absorption (e.g., normalized difference vegetation index) and photosynthetic light‐use efficiency (e.g., photochemical reflectance index). Such sampling should improve our understanding of controls on photosynthetic carbon uptake in diverse tropical forest stands.     

The contribution of cacao agroforests to the conservation of lower canopy ant and beetle diversity in Indonesia

The ongoing destruction of tropical rainforests has increased the interest in the potential value of tropical agroforests for the conservation of biodiversity. Traditional, shaded agroforests may support high levels of biodiversity, for some groups even approaching that of undisturbed tropical forests. However, it is unclear to what extent forest fauna is represented in this diversity and how management affects forest fauna in agroforests. We studied lower canopy ant and beetle fauna in cacao agroforests and forests in Central Sulawesi, Indonesia, a region dominated by cacao agroforestry. We compared ant and beetle species richness and composition in forests and cacao agroforests and studied the impact of two aspects of management intensification (the decrease in shade tree diversity and in shade canopy cover) on ant and beetle diversity. The agroforests had three types of shade that represented a decrease in tree diversity (high, intermediate and low diversity). Species richness of ants and beetles in the canopies of the cacao trees was similar to that found in lower canopy forest trees. However, the composition of ant and beetle communities differed greatly between the agroforest and forest sites. Forest beetles suffered profoundly from the conversion to agroforests: only 12.5% of the beetle species recorded in the forest sites were also found in the agroforests and those species made up only 5% of all beetles collected from cacao. In contrast, forest ants were well represented in agroforests, with 75% of all species encountered in the forest sites also occurring on cacao. The reduction of shade tree diversity had no negative effect on ants and beetles on cacao trees. Beetle abundances and non-forest ant species richness even increased with decreasing shade tree diversity. Thinning of the shade canopy was related to a decrease in richness of forest ant species on cacao trees but not of beetles. The contrasting responses of ants and beetles to shade tree management emphasize that conservation plans that focus on one taxonomic group may not work for others. Overall ant and beetle diversity can remain high in shaded agroforests but the conservation of forest ants and beetles in particular depends primarily on the protection of natural forests, which for forest ants can be complemented by the conservation of adjacent shaded cacao agroforests.     

Imaging spectroscopy links aspen genotype with below-ground processes at landscape scales

Fine-scale biodiversity is increasingly recognized as important to ecosystem-level processes. Remote sensing technologies have great potential to estimate both biodiversity and ecosystem function over large spatial scales. Here, we demonstrate the capacity of imaging spectroscopy to discriminate among genotypes of Populus tremuloides (trembling aspen), one of the most genetically diverse and widespread forest species in North America. We combine imaging spectroscopy (AVIRIS) data with genetic, phytochemical, microbial and biogeochemical data to determine how intraspecific plant genetic variation influences below-ground processes at landscape scales. We demonstrate that both canopy chemistry and below-ground processes vary over large spatial scales (continental) according to aspen genotype. Imaging spectrometer data distinguish aspen genotypes through variation in canopy spectral signature. In addition, foliar spectral variation correlates well with variation in canopy chemistry, especially condensed tannins. Variation in aspen canopy chemistry, in turn, is correlated with variation in below-ground processes. Variation in spectra also correlates well with variation in soil traits. These findings indicate that forest tree species can create spatial mosaics of ecosystem functioning across large spatial scales and that these patterns can be quantified via remote sensing techniques. Moreover, they demonstrate the utility of using optical properties as proxies for fine-scale measurements of biodiversity over large spatial scales.     

Functional diversity of epiphytes in two tropical lowland rainforests, French Guiana: using bryophyte life-forms to detect areas of high biodiversity

Recent studies have described a new tropical lowland forest type in the Guianas, the tropical lowland cloud forest. It is characterized by an enriched epiphytic species diversity particularly for bryophytes compared to common lowland rainforest, and is facilitated by frequent early morning fog events in valley locations. While the increase in epiphytic species diversity in lowland cloud forests has been documented, uncertainties remain as to (1) how this small scale variation in water supply is shaping the functional diversity of epiphytic components in lowland forests, and (2) whether information on functional group composition of epiphytes might aid in discerning these cloud forests from the common lowland rainforest. We compare the distribution of functional groups of epiphytes across height zones in lowland cloud forest and lowland rain forest of French Guiana in terms of biomass, cover as well as the composition of bryophyte life-forms. Both forests differed in functional composition of epiphytes in the canopy, in particular in the mid and outer canopy, with the cloud forest having a higher biomass and cover of bryophytes and vascular epiphytes as well as a richer bryophyte life-form composition. Bryophyte life-forms characteristic for cloud forests such as tail, weft and pendants were almost lacking in the canopies of common rain forest whereas they were frequent in lowland cloud forests. We suggest that ground-based evaluation of bryophyte life-form composition is a straightforward approach for identifying lowland cloud forest areas for conservation, which represent biodiversity hotspots in tropical lowland forests.