Beta-diversity in seasonally dry tropical forests (SDTF) in the Caatinga Biogeographic Domain, Brazil, and its implications for conservation

Tropical biomes are species rich, but some biomes such as seasonally dry tropical forests (SDTFs) are still inadequately studied compared to their co-occurring rain forest and savanna. SDTFs occur in areas of high environmental heterogeneity, resulting in high beta (β)-diversity or species turnover, but this has so far only been accessed using a single β-diversity measure, and at a spatial scale that is of limited applicability for reserve planning. The Caatinga Biogeographic Domain in Brazil contains the largest known extent of SDTF which are poorly studied and inadequately reserved. We therefore studied the variation in species richness and species turnover among SDTF between localities and between known floristic communities. From six localities within the Caatinga Biogeographic Domain we recorded all tree species with a circumference at breast height equaling or exceeding 10 cm within 106 400 m² survey plots. From the species presence/absence data we calculated three measures of β-diversity between pairs of study localities and between different floristic communities representing: (i) species similarity, (ii) differences between species richness, and (iii) species gain and loss. Our results confirm the high β-diversity of SDTFs and species turnover between localities and also between floristic communities. The three indices were also complementary to each other and can be used to maximize accuracy in β-diversity studies. The implications of our study for conservation and reserve planning of SDTFs are discussed.     

The intermediate disturbance hypothesis applies to tropical forests, but disturbance contributes little to tree diversity

The intermediate disturbance hypothesis (IDH) predicts local species diversity to be maximal at an intermediate level of disturbance. Developed to explain species maintenance and diversity patterns in species-rich ecosystems such as tropical forests, tests of IDH in tropical forest remain scarce, small-scale and contentious. We use an unprecedented large-scale dataset (2504 one-hectare plots and 331 567 trees) to examine whether IDH explains tree diversity variation within wet, moist and dry tropical forests, and we analyse the underlying mechanism by determining responses within functional species groups. We find that disturbance explains more variation in diversity of dry than wet tropical forests. Pioneer species numbers increase with disturbance, shade-tolerant species decrease and intermediate species are indifferent. While diversity indeed peaks at intermediate disturbance levels little variation is explained outside dry forests, and disturbance is less important for species richness patterns in wet tropical rain forests than previously thought.     

Floristic diversity of sabal palmetto woodland: an endemic and endangered vegetation type from Mexico

The sabal palmetto woodland is a tropical plant formation dominated by Sabal mexicana, with restricted distribution to southeast Mexico. Sabal palms grow on poor soils but accumulate large quantities of organic substrate in their crowns, harboring a contingent of plants that use it as phorophyte. Although it is a threatened formation, basic information on its biodiversity is scant. We examined the floristic diversity of recruited (diameter at breast height, DBH, ≥1 cm) and understory (DBH ≤ 1 cm) plants, and its variation with anthropogenic disturbance. We also examined the floristic diversity of plants that use the sabal palms as phorophytes, and assessed its variation with human impact. All plants present in transects within a conserved and an adjacent perturbed area were sampled. The list of observed taxa shows that this vegetation has a clear affinity with tropical dry and wet forests of Mexico, with a small representation of taxa from desert ecosystems. The floristic contingent included a total of 81 species in 2000 m². Richness, composition and diversity were affected by disturbance. Recruited and understory vegetation in the disturbed site were 5- and 1.6-times less diverse than in the conserved site, and species of mature, conserved vegetation were substituted by heliophytes in the disturbed site. In contrast, abundance of palms and diversity and identity of epiphytic/hemiepiphytic plants were not affected by disturbance. We show that even monodominated tropical ecosystems growing on poor soils have a high floristic diversity and that current anthropogenic impact threatens not only species and populations but also entire plant formations.     

A transitional xerophytic tropical plant community of the Cape Region,Baja California

Abstract. Our main objective was to use comparative floristic and structural criteria in order to understand the differences in assemblages brought about by the meeting of arid and dry‐tropical environments. We sampled 33 vegetation plots and recorded all perennial species. The data were subjected to multivariate analysis, including Principal Component Analysis (PCA). The floristic variation detected by the PCA was strongly related to altitude and rainfall. The gradient detected by the ordination procedure had Tropical deciduous forest (TDF) and Sonoran Desert (SD) at opposite extremes and Cape sarcocaulescent shrubland (CSCS) in an intermediate position. The numerical classification algorithm detected six distinct groups of species with clearly identifiable field distributions. Vegetation structure also differed significantly between the regions. SD had the lowest species‐richness (α‐diversity) and inter‐site similarity in the CSCS was highest, indicating that this environment is relatively uniform and has low species turnover (β‐diversity). It is concluded that the CSCS is distinctly different from the sarcocaulescent shrublands of the more northern plant communities of the peninsula, where Sonoran Desert floristic affinities prevail. The floristic composition of the CSCS is rich in dry‐tropical affinities, its species richness is higher, it is homogeneous in its species content and turnover and is more dense with a taller canopy than the northern desert scrubs. It is proposed that the boundary line separating the Cape sediments of granitic origin from the basalt‐derived sediments of the northern‐lying Sierra de la Giganta should be used as an easily identifiable landscape trait to delimitate this unique community.     

Rainfall seasonality and drought performance shape the distribution of tropical tree species in Ghana

Tree species distribution in lowland tropical forests is strongly associated with rainfall amount and distribution. Not only plant water availability, but also irradiance, soil fertility, and pest pressure covary along rainfall gradients. To assess the role of water availability in shaping species distribution, we carried out a reciprocal transplanting experiment in gaps in a dry and a wet forest site in Ghana, using 2,670 seedlings of 23 tree species belonging to three contrasting rainfall distributions groups (dry species, ubiquitous species, and wet species). We evaluated seasonal patterns in climatic conditions, seedling physiology and performance (survival and growth) over a 2‐year period and related seedling performance to species distribution along Ghana's rainfall gradient. The dry forest site had, compared to the wet forest, higher irradiance, and soil nutrient availability and experienced stronger atmospheric drought (2.0 vs. 0.6 kPa vapor pressure deficit) and reduced soil water potential (?5.0 vs. ?0.6 MPa soil water potential) during the dry season. In both forests, dry species showed significantly higher stomatal conductance and lower leaf water potential, than wet species, and in the dry forest, dry species also realized higher drought survival and growth rate than wet species. Dry species are therefore more drought tolerant, and unlike the wet forest species, they achieve a home advantage. Species drought performance in the dry forest relative to the wet forest significantly predicted species position on the rainfall gradient in Ghana, indicating that the ability to grow and survive better in dry forests and during dry seasons may allow species to occur in low rainfall areas. Drought is therefore an important environmental filter that influences forest composition and dynamics. Currently, many tropical forests experience increase in frequency and intensity of droughts, and our results suggest that this may lead to reduction in tree productivity and shifts in species distribution.     

High butterfly beta diversity between Brazilian cerrado and cerrado–caatinga transition zones

Tropical dry forests are among the most diverse and threatened habitats in the world, yet they are rarely protected and remain poorly studied. In Brazil, dry forests are naturally fragmented and embedded within various biomes, thus making it important to assess biotic homogeneity among geographically separated forest fragments. We sampled 7732 individuals belonging to 48 species to quantify the diversity of fruit-feeding butterfly communities at four Brazilian dry forest sites, and found differences in community structure between northern and central sites. Species richness per plot was the same in both areas, but abundance per plot was higher in northern sites. Species composition differed between sites mostly due to species of Satyrinae. Additive partitioning showed that beta diversity corresponded to 70.1% of all diversity. Rather than species loss, beta diversity primarily represented species turnover that was potentially driven by differences in the surrounding habitats. Butterfly community composition and abundance were influenced by vegetation where abundance increased with tree density and basal area, and decreased with the average tree height. Butterfly species richness and abundance were higher in the wet season than in the dry season, and all species sampled in the dry season were present in the wet season. Differences in community structure across relatively short geographic distances in the same type of habitat highlight the importance of conserving tropical dry forest fragments to ensure the maintenance of butterfly diversity and, presumably, other insects.     

Termite diversity in Neotropical dry forests of Colombia and the potential role of rainfall in structuring termite diversity

Termites are ecosystem engineers that play an important role in the biotransformation and re‐distribution of nutrients in soil. The dry forests are endemic repositories, but at same time, they are most threatened by extensive livestock and crop farming, fires, and climate change. In Colombia, the best‐protected dry forests are located in the north. The termite fauna of dry forests are poorly known. The aim was to identify the termite species occurring in tropical dry forests of the Colombian Caribbean coast in relation to diet and precipitation, temperature, elevation, and soil properties. A total of 32 species in 1,103 occurrences were found. Termitidae accounted for 78% of the species richness with the Anoplotermes‐group, Microcerotermes, and Nasutitermes being the dominant genera. Differences in species composition and abundance were found across sites. These differences may be linked to anthropogenic disturbance and polygyny and polydomy. Strikingly, our highest elevation site (334 m) had the highest species richness much higher than the two lower elevation sites. This implies an inversion of the common elevation‐diversity gradient, also found for termites which can be explained by increasing precipitation with elevation in the dry forest. An analysis of termite species richness at the global scale confirms that termite species richness correlates positively with rainfall. Hence, rainfall seems to positively affect termite diversity. In line, the studied Colombian tropical dry forests had low diversity compared to rain forests. A decline of species‐rich soil‐feeding termites with increasing aridity may explain why the highest termite diversity occurs in humid tropical rain forests. Abstract in Spanish is available with online material.     

Avian Composition Co‐varies with Floristic Composition and Soil Nutrient Concentration in Amazonian Upland Forests

Spatial heterogeneity in the plant species composition of tropical forests is expected to influence animal species abundance and composition because vegetation constitutes the primary habitat feature for forest animals. Floristic variation is tied to variation in soils, so edaphic properties should ultimately influence animal species composition as well. The study of covariation in floristic and faunistic turnover has been hindered by the difficulty of completing coordinated surveys in hyperdiverse tropical communities, but this can be overcome with the use of a few plant taxa that function as surrogates for general floristic turnover. We used avian and plant transect surveys and soil sampling in a western Amazonian upland (terra firme) forest landscape to test whether spatial variation in bird community composition is associated with floristic turnover and corresponding edaphic gradients. Partial Mantel tests and Non‐metric Multidimensional Scaling showed floristic distinctiveness between two forest types closely associated with differences in soil cation concentrations, and differences in both floristic composition and cation concentrations were further linked to compositional differences in avian species, independent of geographic distances among sites. Ten percent of bird species included in Indicator Species Analyses showed significant associations with one of the two forest types. The upland forest types that we sampled, each corresponding to a different geological formation, are intermediate relative to edaphically extreme environments in the region. Models of avian diversification should take into account this environmental heterogeneity, as should conservation planning approaches that aim to represent faunal diversity. Abstract in Spanish is available in the online version of this article.     

Vegetation structure determines insect herbivore diversity in seasonally dry tropical forests

Vegetation structure can often determine insect herbivore fauna in forests, but this mechanism has been demonstrated in seasonally dry tropical forests (SDTFs) only at small spatial scales. In this study we evaluated the effects of the geographical location of SDTFs and vegetation structure on insect herbivore communities (leaf-chewing and sap-sucking guilds) in three Brazilian ecoregions (Cerrado, Cerrado/Caatinga transition, and Caatinga). We tested the following predictions: (1) insect herbivore species composition, richness, abundance and beta diversity differ among forests in different ecoregions; (2) insect richness, abundance and beta diversity are positively related to tree richness and density; (3) spatial turnover of species is the primary mechanism that generates herbivorous insect β-diversity in different ecoregions, and is positively influenced by tree richness. The composition, richness, and abundance of herbivorous insects differed over SDFs along the gradient of Cerrado and Caatinga. Both herbivore guilds responded positively to tree richness. Tree density only determined the richness and abundance of sap-sucking herbivores. Insect β-diversity was similar among Cerrado and transition areas, but lower in Caatinga itself; β-diversity was also positively affected by tree richness. Species turnover, as opposed to nestedness, was the main mechanism generating β-diversity, but itself was not related to tree richness. We demonstrate in this study the importance of landscape diversity and availability of local resources for herbivorous insect communities, and we emphasize the importance of SDTF conservation in different ecoregions as a result of species turnover.     

Phyllostomid Bat Community Structure and Abundance in Two Contrasting Tropical Dry Forests1

Although tropical wet forests are generally more diverse than dry forests for many faunal groups, few studies have compared bat diversity among dry forests. I compared ground level phyllostomid bat community structure between two tropical dry forests with different precipitation regimes. Parque National Palo Verde in northwestern Costa Rica represents one of the wettest tropical dry forests (rainfall 1.5 m/yr), whereas the Chamela‐Cuixmala Biosphere Reserve on the Pacific coast of central Mexico represents one of the driest (750 mm/yr). Mist net sampling was conducted at the two study sites to compare changes in ground level phyllostomid bat community structure between regions and seasons. Palo Verde was more diverse than Chamela and phyllostomid species showed low similarity between sites (Classic Jaccard = 0.263). The distinct phyllostomid communities observed at these two dry forest sites demonstrates that variants of tropical dry forest can be sufficiently different in structure and composition to affect phyllostomid communities. At both dry forest sites, abundance of the two most common foraging guilds (frugivores and nectarivores) differed between seasons, with greatest numbers of individuals captured coinciding with highest chiropterophilic resource abundance.     

Changes in tree phenology along natural regeneration in a seasonally dry tropical forest

The applicability of succession models from temperate and tropical wet forests to threatened seasonally dry tropical forests (SDTFs) is questioned. Plant phenology affects ecosystem functions and changes along forest regeneration gradient. To investigate the recovery of ecological functions after disturbances in a SDTF, we recorded the vegetative and reproductive phenologies for trees (DBH >5 cm) for 17 months in southeast Brazil in three successional stages: early (10–15 years after clearing), intermediate (25–30) and late (>50). The vegetative phenology of the 523 individuals was strongly seasonal, with 3% of individuals presenting green leaves in a deciduous dry season. Besides structural and floristic differences, phenological trends were similar between the later stages. Reproduction occurred with higher intensities in the early stage and in the advanced stages only in the dry season, providing key resources to local fauna. The studied SDTF is resilient to ecological functions, rapidly recovering functional processes. The integration of structural and functional knowledge of succession of STDFs may lead to better management of its secondary remnants. Our study suggests that classical forest succession theory developed for other ecosystems may not fully reflect the pattern of SDTF succession, an ecosystem that originally covered 42% of the earth's tropical and subtropical landmass.     

Arbuscular mycorrhizal communities in tropical forests are affected by host tree species and environment

Arbuscular mycorrhizal (AM) fungi are mutualists with plant roots that are proposed to enhance plant community diversity. Models indicate that AM fungal communities could maintain plant diversity in forests if functionally different communities are spatially separated. In this study we assess the spatial and temporal distribution of the AM fungal community in a wet tropical rainforest in Costa Rica. We test whether distinct fungal communities correlate with variation in tree life history characteristics, with host tree species, and the relative importance of soil type, seasonality and rainfall. Host tree species differ in their associated AM fungal communities, but differences in the AM community between hosts could not be generalized over life history groupings of hosts. Changes in the relative abundance of a few common AM fungal species were the cause of differences in AM fungal communities for different host tree species instead of differences in the presence and absence of AM fungal species. Thus, AM fungal communities are spatially distinguishable in the forest, even though all species are widespread. Soil fertility ranging between 5 and 9 Mg/ha phosphorus did not affect composition of AM fungal communities, although sporulation was more abundant in lower fertility soils. Sampling soils over seasons revealed that some AM fungal species sporulate profusely in the dry season compared to the rainy season. On one host tree species sampled at two sites with vastly different rainfall, relative abundance of spores from Acaulospora was lower and that of Glomus was relatively higher at the site with lower and more seasonal rainfall.     

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.     

Host Specificity, Alpha- and Beta-Diversity of Phytophagous Beetles in Two Tropical Forests in Panama

Species diversity, host specificity and species turnover among phytophagous beetles were studied in the canopy of two tropical lowland forests in Panama with the use of canopy cranes. A sharp rainfall gradient occurs between the two sites located 80 km apart. The wetter forest is located in San Lorenzo Protected Area on the Caribbean side of the isthmus, and the drier forest is a part of the Parque Natural Metropolitano close to Panama City on the Pacific slope. Host specificity was measured as effective specialization and recorded by probability methods based on abundance categories and feeding records from a total of 102 species of trees and lianas equally distributed between the two sites. The total material collected included more than 65,000 beetles of 2462 species, of which 306 species were shared between the two sites. The wet forest was 37% more species rich than the dry forest due to more saproxylic species and flower visitors. Saproxylic species and flower visitors were also more host-specific in the wet forest. Leaf chewers showed similar levels of species richness and host specificity in both forests. The effective number of specialized species per plant species was higher in the wet forest. Higher levels of local alpha- and beta-diversity as well as host specificity based on present data from a tropical wet forest, suggests higher number of species at regional levels, a result that may have consequences for ecological estimates of global species richness.     

Domatia morphology and mite occupancy of Psychotria horizontalis (Rubiaceae) across the Isthmus of Panama

Leaf domatia are small plant structures in vein axials on the undersides of leaves that are often inhabited by mites of several species. The mites are presumed to benefit the plant because they are predatory or fungivorous. The domatia are thought to provide the mites shelter from predators and changes in relative humidity, and in exchange, the mites protect the plant from small herbivores and fungal spores. Differences in relative humidity can affect food availability, changing the interaction between plants and mites. We examined domatium morphology of the shrub Psychotria horizontalis (Rubiaceae) and its associated mite diversity at three sites along the rainfall gradient of the Isthmus of Panama, during the dry and wet seasons. The dry forest had a domatium morphology consistent with providing greater desiccation protection, with trichomes and a smaller domatium opening relative to domatium size (size/opening ratio). Additionally, this size/opening ratio was significantly higher in the dry season than in the wet season at all three sites. Mite diversity was highest at the intermediate rainfall site with a large degree of overlap with the other sites, whereas the dry site and wet site shared few mite species. More fungivorous mites were present in the moist forests and more facultative feeders on fungal spores and small mites in the dry forest. The average mite size at each site matched the average domatium size at each site. The dry forest had small mites in small domatia, whereas the moist forests had larger mites in larger domatia. While these data are primarily observational, the site and seasonal differences in domatium morphology and mite diversity are consistent with two main hypotheses: (1) that protection from changes in humidity would be particularly important when humidity was low, such as in the dry forest and during the dry season (2) more fungivorous mites would be found in domatia of the moist forests. The data presented here further highlight the close adaptive relationship between leaf domatia on plants and the mites that inhabit them.     

Effects of Drought,Pest Pressure and Light Availability on Seedling Establishment and Growth: Their Role for Distribution of Tree Species across a Tropical Rainfall Gradient

Tree species distributions associated with rainfall are among the most prominent patterns in tropical forests. Understanding the mechanisms shaping these patterns is important to project impacts of global climate change on tree distributions and diversity in the tropics. Beside direct effects of water availability, additional factors co-varying with rainfall have been hypothesized to play an important role, including pest pressure and light availability. While low water availability is expected to exclude drought-intolerant wet forest species from drier forests (physiological tolerance hypothesis), high pest pressure or low light availability are hypothesized to exclude dry forest species from wetter forests (pest pressure gradient and light availability hypothesis, respectively). To test these hypotheses at the seed-to-seedling transition, the potentially most critical stage for species discrimination, we conducted a reciprocal transplant experiment combined with a pest exclosure treatment at a wet and a dry forest site in Panama with seeds of 26 species with contrasting origin. Establishment success after one year did not reflect species distribution patterns. However, in the wet forest, wet origin species had a home advantage over dry forest species through higher growth rates. At the same time, drought limited survival of wet origin species in the dry forest, supporting the physiological tolerance hypothesis. Together these processes sort species over longer time frames, and exclude species outside their respective home range. Although we found pronounced effects of pests and some effects of light availability on the seedlings, they did not corroborate the pest pressure nor light availability hypotheses at the seed-to-seedling transition. Our results underline that changes in water availability due to climate change will have direct consequences on tree regeneration and distributions along tropical rainfall gradients, while indirect effects of light and pests are less important.     

Epiphytic orchids in tropical dry forests of Yucatan,Mexico – Species occurrence,abundance and correlations with host tree characteristics and environmental conditions

Tropical dry forests have been less studied in terms of their resident epiphyte flora compared to wet forests. We studied five species of epiphytic orchids in two dry forest fragments differing in tree composition, stature and rainfall regime. We compared the vertical distribution within the host tree, epiphyte-host associations and seasonal variation in microclimatic conditions in a tropical dry deciduous (Celestún) and a semi-deciduous forest (Kaxil-Kiuic) of the Yucatan Peninsula, Mexico, during the wet, early dry and dry seasons. Light, vapor pressure deficit, air temperature, and dew were measured on two heights (1.5 and 3.5 m) of the host with the highest abundance of orchids. Surprisingly, orchid abundance was higher in the Celestún deciduous forest, the site with low precipitation. High epiphyte abundance in the middle canopy stratum of the hosts in both forests was arguably related to a favorable combination of micro-environmental factors. In both forests, about 90% of all orchids grew on a single host tree species. Although bark roughness and the area of the substrate were the most important host characteristics that influenced the abundance of orchids in the Celestún deciduous forest, this did not explain this preference. Climatic variation was greater among seasons than between microenvironments in the host trees. The most abundant species, Encyclia nematocaulon, had a great capacity to occupy different strata in both forests, even in tree tops with very large micro-environmental fluctuations during the year.     

Are long‐unburnt eucalypt forest patches important for the conservation of plant species diversity?

Question: Are long‐unburnt patches of eucalypt forest important for maintaining floristic diversity? Location: Eucalyptus forests of southeastern New South Wales, Australia. Methods: Data from 976 sites representing a range of fire history from three major vegetation formations – shrubby dry sclerophyll forest (SF), grassy dry SF and wet SF – were analysed. Generalized linear models were used to examine changes in species richness with increasing time since wildfire and analysis of similarities to examine changes in community composition. Chi‐squared tests were conducted to examine the distribution of individual species across four time since fire categories. Results: Plant species relationships to fire varied between the three formations. Shrubby dry SF supported lower plant species richness with increasing time since wildfire and this was associated with shifts in community composition. Grassy dry SF showed significant shifts in community composition and species richness in relation to time, with a peak in plant species richness 20–30 yr post fire (either prescribed fire or wildfire). Wet SF increased in species richness until 10–20 yr post wildfire then displayed a general declining trend. Species richness in each vegetation type was not related to the fire frequencies and fire intervals observed in this study. Conclusions: Long‐unburnt (30–50 yr post wildfire) forests appeared to play a minor role in the maintenance of plant species diversity in dry forest systems, although this was more significant in wet forests. Maintenance of a range of fire ages within each vegetation formation will assist in maintaining floristic diversity within regions.     

Prevalence of Tree Regeneration by Sprouting and Seeding Along a Rainfall Gradient in Hawai'i

Drought stress in tropical dry forests is thought to result in greater asexual regeneration via vegetative sprouting ( e.g ., basal, root, and branch layering) than occurs in moister tropical forests. We tested this hypothesis by examining the prevalence of tree sprouting and seeding in tropical forests located along a rainfall gradient on the island of Hawai'i. Additionally, we examined the potential for novel disturbance, feral pig Sus scrofa rooting and trampling, to alter patterns in tree regeneration mode. We found greater sprouting (in terms of relative density and basal area) in dry forests than in mesic and wet forests, supporting the hypothesis. We also found that feral pig disturbance is negatively correlated with the relative density and basal area of seedlings in wet forests, but is positively correlated with the relative importance of sprouting, and the richness and diversity of sprouting species. Our results suggest rainfall regimes may be an important factor controlling broad-scale patterns in tree regeneration mode, and that exotic ungulates can significantly modify such patterns with potential consequences for the structure and dynamics of tree populations and communities.     

Quantifying the deciduousness of tropical forest canopies under varying climates

Abstract. Deciduousness is an important functional attribute of tropical trees, reflecting climatic conditions. Precisely quantifying and mapping deciduousness in tropical forests will be necessary for calibrating remote sensing images which attempt to assess canopy properties such as carbon cycling, productivity, or chlorophyll content. We thus set out to assess the degree of canopy deciduousness in three moist, semi‐deciduous tropical forests in central Panama. One site is a 6‐ha research plot near the Atlantic coast of Panama, where rainfall is 2830 mm/yr. The second site is a 50‐ha plot on Barro Colorado Island, near the center of the isthmus of Panama, where rainfall is 2570 mm/yr, and the final site is a 4‐ha plot near the Pacific coast of Panama, where rainfall is 2060 mm/yr. At each site, a random sample of trees from all canopy species (those with individuals ≥ 30 cm DBH) were visited and scored for deciduousness three times during the 1997 dry season. The estimated peak fraction of deciduous individuals in the canopy at the wetter site was 4.8%, at the intermediate site, 6.3%, and at the drier site, 24.3%. The estimated fraction of crown area deciduous peaked at 3.6%, 9.7%, and 19.1% at the wet, medium, and dry sites respectively. The percentage of canopy species that was deciduous –14%, 28%, and 41%–was much higher than the percentage of deciduous individuals, because not all individuals of deciduous species were deciduous. During the 1999 dry season, every individual of all the deciduous species was visited at the two drier sites, and the total number of deciduous trees observed closely matched the estimated numbers based on the smaller 1997 samples.