Diversity of dry forest epiphytes along a gradient of human disturbance in the tropical Andes

Question: Disturbance effects on dry forest epiphytes are poorly known. How are epiphytic assemblages affected by different degrees of human disturbance, and what are the driving forces? Location: An inter‐Andean dry forest landscape at 2300 m elevation in northern Ecuador. Methods: We sampled epiphytic bryophytes and vascular plants on 100 trees of Acacia macracantha in five habitats: closed‐canopy mixed and pure acacia forest (old secondary), forest edge, young semi‐closed secondary woodland, and isolated trees in grassland. Results: Total species richness in forest edge habitats and on isolated trees was significantly lower than in closed forest types. Species density of vascular epiphytes (species per tree) did not differ significantly between habitat types. Species density of bryophytes, in contrast, was significantly lower in edge habitat and on isolated trees than in closed forest. Forest edge showed greater impoverishment than semi‐closed woodland and similar floristic affinity to isolated trees and to closed forest types. Assemblages were significantly nested; habitat types with major disturbance held only subsets of the closed forest assemblages, indicating a gradual reduction in niche availability. Distance to forest had no effect on species density of epiphytes on isolated trees, but species density was closely correlated with crown closure, a measure of canopy integrity. Main conclusions: Microclimatic changes but not dispersal constraints were key determinants of epiphyte assemblages following disturbance. Epiphytic cryptogams are sensitive indicators of microclimate and human disturbance in montane dry forests. The substantial impoverishment of edge habitat underlines the need for fragmentation studies on epiphytes elsewhere in the Tropics.     

Habitat isolation changes the beta diversity of the vascular epiphyte community in lower montane forest,Veracruz, Mexico

Habitat isolation is one of the most important factors endangering the biodiversity, but little research has been done with vascular epiphytes. In order to understand the effect of isolation on the epiphyte community, we studied epiphyte diversity on three plots in a forest fragment, two riparian forest fragments, and in isolated pastureland trees. We found 118 vascular epiphyte species. On forest plots, both epiphyte richness per tree (S_tree) and species turnover rate within trees (β_tree) registered the highest values, although the lowest S_tree diversity was also found there; additionally inside the forest were host species with clearly different epiphyte community. S_tree and β_tree diversities of riparian fragments behaved similarly to those of the forest. Isolated trees had the second highest S_tree diversity, although their β_tree diversity was the lowest. In the forest plots were both, the highest and lowest expected accumulated richness (α diversity); on riparian fragments it was intermediate, and the second lowest α diversity was registered for isolated trees. Species turnover rate among plots (β) was high and was associated with both, isolation and a distance gradient from permanent water sources. The epiphyte community on isolated trees was clearly different to the other habitats. Results suggest that deforestation eliminated dry areas and specific hosts that were important for the maintenance of epiphyte species richness. In pastureland trees the epiphyte β_tree diversity diminished, suggesting a simplification of the environment for epiphytes and causing a low α diversity.     

Structure of the Epiphyte Community in a Tropical Montane Forest in SW China

Vascular epiphytes are an understudied and particularly important component of tropical forest ecosystems. However, owing to the difficulties of access, little is known about the properties of epiphyte-host tree communities and the factors structuring them, especially in Asia. We investigated factors structuring the vascular epiphyte-host community and its network properties in a tropical montane forest in Xishuangbanna, SW China. Vascular epiphytes were surveyed in six plots located in mature forests. Six host and four micro-site environmental factors were investigated. Epiphyte diversity was strongly correlated with host size (DBH, diameter at breast height), while within hosts the highest epiphyte diversity was in the middle canopy and epiphyte diversity was significantly higher in sites with canopy soil or a moss mat than on bare bark. DBH, elevation and stem height explained 22% of the total variation in the epiphyte species assemblage among hosts, and DBH was the most important factor which alone explained 6% of the variation. Within hosts, 51% of the variation in epiphyte assemblage composition was explained by canopy position and substrate, and the most important single factor was substrate which accounted for 16% of the variation. Analysis of network properties indicated that the epiphyte host community was highly nested, with a low level of epiphyte specialization, and an almost even interaction strength between epiphytes and host trees. Together, these results indicate that large trees harbor a substantial proportion of the epiphyte community in this forest.     

海南岛热带天然针叶林附生维管植物多样性和分布

作为热带林中一个重要的特征性组分, 附生维管植物对于维持热带森林的物种多样性及其生态系统功能均具有重要作用。该文首次系统地报道了热带天然针叶林中的附生维管植物多样性和分布特征。以海南岛霸王岭国家级自然保护区保存完好的热带天然针叶林(我国唯一较大面积分布的南亚松(Pinus latteri)天然林)中的附生维管植物为研究对象, 通过样带调查(共设置12个10 m × 50 m的样带, 记录每个样带内胸径(DBH) ≥ 5 cm树木上附生维管植物的物种名称、株数及附生高度), 分析附生维管植物的物种多样性和空间分布特征。结果表明: 1)热带针叶林0.6 hm ²面积内共有附生维管植物769株, 分属于7科17属27种, 附生兰科植物和萝摩科植物为优势类群; 2)附生维管植物在水平方向上呈现出聚集分布; 3)附生维管植物在垂直方向上, 在中等高度层次(10-20 m)分布最多, 在下层(0-5 m)也有较多的分布; 4)少数附生维管植物对南亚松表现出一定的选择性, 如华南马尾杉(Phlegmariurus fordii)、玫瑰毛兰(Eria rosea)、眼树莲(Dischidia chinensis)和铁草鞋(Hoya pottsii)等; 5)附生维管植物的物种丰富度及多度与宿主胸径均存在显著的正相关关系。     

Long‐term changes of the vascular epiphyte assemblage on the palm Socratea exorrhiza in a lowland forest in Panama

Question: What are the qualitative and quantitative long‐term changes in the vascular epiphyte assemblage on a particular host tree species? Location: Lowland rain forest of the San Lorenzo Crane Plot, Republic of Panama. Methods: We followed the fate of the vascular epiphyte assemblage on 99 individuals of the palm Socratea exorrhiza by three censuses over the course of five years. Results: The composition of the epiphyte assemblage changed little during the course of the study. While the similarity of epiphyte vegetation decreased on individual palms through time, the similarity analysed over all palms increased. Even well established epiphyte individuals experienced high mortality with only 46% of the originally mapped individuals surviving the following five years. We found a positive correlation between host tree size and epiphyte richness and detected higher colonization rates of epiphytes per surface area on larger trees. Conclusions Epiphyte assemblages on individual S. exorrhiza trees were highly dynamic while the overall composition of the epiphyte vegetation on the host tree species in the study plot was stable. We suggest that higher recruitment rates, due to localized seed dispersal by already established epiphytes, on larger palms promote the colonization of epiphytes on larger palms. Given the known growth rates and mortality rates of the host tree species, the maximum time available for colonization and reproduction of epiphytes on a given tree is estimated to be ca. 60 years. This time frame will probably be too short to allow assemblages to be ever saturated.     

Can spatial variation in epiphyte diversity and community structure be predicted from sampling vascular epiphytes alone?

Aim Non‐vascular epiphytes have been largely ignored in studies examining the biotic and abiotic determinants of spatial variation in epiphyte diversity. Our aim was to test whether the spatial patterning of species richness, biomass and community composition across geographic regions, among trees within regions, and among branches within trees is consistent between the vascular and non‐vascular components of the temperate rain forest flora. Location Coastal lowland podocarp‐broadleaved forests on the west coast of the South Island of New Zealand. Methods We collected single samples (30 × 25 cm) from 96 epiphyte assemblages located on the inner branches of 40 northern rata (Metrosideros robusta) trees. For each sample, branch characteristics such as branch height, branch diameter, branch angle, branch aspect, and minimum and maximum epiphyte mat depth were recorded. The biomass for each individual epiphyte species was determined. Results Northern rata was host to a total of 157 species, comprising 32 vascular and 125 non‐vascular species, with liverworts representing 41% of all species. Within epiphyte mats, the average total organic biomass of 3.5 kg m^?2 of branch surface area consisted largely of non‐living biomass and roots. Vascular and non‐vascular epiphytes showed strikingly different spatial patterns in species richness, biomass and composition between sites, among trees within sites, and among branches within trees, which could not be explained by the branch structural characteristics we measured. The two plant groups had no significant association in community composition (r = 0.04, P = 0.08). However, the species richness of vascular plant seedlings was strongly linked to the presence/absence of lichens. Main conclusions Non‐vascular plants contributed substantially to the high species richness and biomass recorded in this study, which was comparable to that of some tropical rain forests. High variability in community composition among epiphyte mats, and very low correlation with any of the environmental factors measured possibly indicate high levels of stochasticity in seed or spore colonization, establishment success or community assembly among branches in these canopy communities. Although we found some evidence that vascular plant seedling establishment was linked to the presence of lichens and the biomass of non‐living components in the epiphyte mats, there was no correlation in the spatial patterning or determinants of species richness between non‐vascular and vascular plants. Consequently, variation in total epiphyte biodiversity could not be predicted from the measurement of vascular plant diversity alone, which highlights the crucial importance of sampling non‐vascular plants when undertaking epiphyte community studies.     

Composition and distribution of vascular epiphytes in a tropical semideciduous forest,Ghana

The composition and distribution of vascular epiphytes were studied in two 1‐ha plots in the KNUST Botanic garden, Ghana. One‐hectare plot each was randomly set up in secondary and cultivated forests for the identification and enumeration of trees and shrubs (≥10 cm dbh), and epiphytes. Each tree was carefully examined, noting the presence, positions and life‐forms of all epiphytes. Twenty‐nine epiphyte (29) species belonging to fourteen genera and eleven families were identified in the study. These were hosted by 48 tree species and occurred in three life‐forms: hemi‐epiphytes (45%), casual epiphytes (45%) and true epiphytes (10%). The vascular epiphyte species made up 25.7% of all the identified plant species (excluding herbs and climbers) encountered. Host species (P < 0.001), habitat (P = 0.001) and their interaction (P < 0.001) had strong effects on epiphyte composition in the forests. Moraceae was the most dominant family (44.8%), while Nephrolepis undulata J. Sm. and N. biserrata (Sw.) Scott. were the commonest species of epiphytes. In terms of vertical distribution, most epiphytes were located on the trunk, while a few occurred in the canopy.     

Diversity patterns of vascular epiphytes along an elevational gradient in the Andes

Aim To document the elevational pattern of epiphyte species richness at the local scale in the tropical Andes with a consistent methodology. Location The northern Bolivian Andes at 350–4000 m above sea level. Methods We surveyed epiphytic vascular plant assemblages in humid forests in (a) single trees located in (b) 90 subplots of 400 m² each located in (c) 14 plots of 1 ha each. The plots were separated by 100–800 m along the elevational gradient. Results We recorded about 800 epiphyte species in total, with up to 83 species found on a single tree. Species richness peaked at c. 1500 m and declined by c. 65% to 350 m and by c. 99% to 4000 m, while forests on mountain ridges had richness values lowered by c. 30% relative to slope forests at the same elevations. The hump‐shaped richness pattern differed from a null‐model of random species distribution within a bounded domain (the mid‐domain effect) as well as from the pattern of mean annual precipitation by a shift of the diversity peak to lower elevations and by a more pronounced decline of species richness at higher elevations. With the exception of Araceae, which declined almost monotonically, all epiphyte taxa showed hump‐shaped curves, albeit with slightly differing shapes. Orchids and pteridophytes were the most species‐rich epiphytic taxa, but their relative contributions shifted with elevation from a predominance of orchids at low elevations to purely fern‐dominated epiphyte assemblages at 4000 m. Within the pteridophytes, the polygrammoid clade was conspicuously overrepresented in dry or cold environments. Orchids, various small groups (Cyclanthaceae, Ericaceae, Melastomataceae, etc.), and Bromeliaceae (below 1000 m) were mostly restricted to the forest canopy, while Araceae and Pteridophyta were well represented in the forest understorey. Main conclusions Our study confirms the hump‐shaped elevational pattern of vascular epiphyte richness, but the causes of this are still poorly understood. We hypothesize that the decline of richness at high elevations is a result of low temperatures, but the mechanism involved is unknown. The taxon‐specific patterns suggest that some taxa have a phylogenetically determined propensity for survival under extreme conditions (low temperatures, low humidity, and low light levels in the forest interior). The three spatial sampling scales show some different patterns, highlighting the influence of the sampling methodology.     

Epiphyte host preferences and host traits: mechanisms for species-specific interactions

We investigated species-specific relationships among two species of vascular epiphytes and ten host tree species in a coastal plain forest in the southeastern United States. The epiphytes Tillandsia usneoides and Polypodium polypodioides were highly associated with particular host species in the field, but host traits that favored colonization were inadequate to fully explain the epiphyte-host associations for either epiphyte. Field transplant experiments that bypassed epiphyte colonization demonstrated that the growth of epiphytes was significantly higher on host tree species that naturally bore high epiphyte loads than on host species with few or no epiphytes. These species-specific relationships were highly correlated with the water-holding capacity of the host tree's bark. Positive and negative effects of throughfall, light attenuation by the canopy, and bark stability did not explain the overall patterns of host specificity, but did correlate with some epiphyte-host species relationships. The relative importance of particular host traits differed between the "atmospheric epiphyte" Tillandsia, and the fern Polypodium, which roots in the bark of its hosts. Species-specific interactions among plants, such as those described here, suggest that communities are more than individualistic assemblages of co-occurring species.     

The nutrient status of epiphytes and their host trees along an elevational gradient in Costa Rica

Vascular epiphytes are a conspicuous and highly diverse group in tropical wet forests; yet, we understand little about their mineral nutrition across sites. In this study, we examined the mineral nutrition of three dominant vascular epiphyte groups: ferns, orchids, and bromeliads, and their host trees from samples collected along a 2600 m elevational gradient in the tropical wet forests of Costa Rica. We predicted that the mineral nutrition of ferns, orchids, and bromeliads would differ because of their putative differences in nutrient acquisition mechanisms and nutrient sources—atmospherically dependent, foliar feeding bromeliads would have lower nitrogen (N) and phosphorous (P) concentrations and more depleted δ¹⁵N values than those in canopy soil-rooted ferns because canopy soil is higher in available N, and more enriched in δ¹⁵N than the atmospheric sources of precipitation and throughfall. We also predicted that epiphyte foliar chemistry would mirror that of host trees because of the likely contribution of host trees to the nutrient cycle of epiphytes via foliar leaching and litter contributions to canopy soil. In the same vein, we predicted that epiphyte and host tree foliar chemistry would vary with elevation reflecting ecosystem-level nutrients—soil N availability increases and P availability decreases with increasing elevation. Our results confirmed that canopy soil-rooted epiphytes had higher N concentrations than atmospheric epiphytes; however, our predictions were not confirmed with respect to P which did not vary among groups indicating fixed P availability within sites. In addition, foliar δ¹⁵N values did not match our prediction in that canopy soil-rooted as well as atmospheric epiphytes had variable signatures. Discriminant function analysis (DFA) on foliar measurements determined that ferns, orchids, and bromeliads are statistically distinct in mineral nutrition. We also found that P concentrations of ferns and orchids, but not bromeliads, were significantly correlated with those of host trees indicating a possible link in their mineral nutrition’s via canopy soil. Interestingly, we did not find any patterns of epiphyte foliar chemistry with elevation. These data indicate that the mineral nutrition of the studied epiphyte groups are distinct and highly variable within sites and the diverse uptake mechanisms of these epiphyte groups enhance resource partitioning which may be a mechanism for species richness maintenance in tropical forest canopies.     

Conservation of Vascular Epiphyte Diversity in Shade Cacao Plantations in the Chocó Region of Ecuador

To assess the contributions of rustic shade cacao plantations to vascular epiphyte conservation, we compared epiphyte species richness, abundance, composition, and vertical distributions on shade trees and in the understories of six plantations and adjacent natural forests. On three phorophytes and three 10 × 10 m understory plots in each of the agroforestry plantations and natural forests, 54 and 77 species were observed, respectively. Individual-based rarefaction curves revealed that epiphyte species richness was significantly higher on forest phorophytes than on cacao farm shade trees; detailed analyses showed that the differences were confined to the inner and outer crown zones of the phorophytes. No differences in epiphyte species richness were found in understories. Araceae, Piperaceae, and Pteridophyta were less species-rich in plantations than in forests, while there were no differences in Orchidaceae and Bromeliaceae. Regression analysis revealed that epiphyte species richness on trunks varied with canopy cover, while abundance was more closely related to soil pH, canopy cover, and phorophyte height. For crown epiphytes, phorophyte diameter at breast height (dbh) explained much of the variation in species richness and abundance. There were also pronounced downward shifts in the vertical distributions of epiphyte species in agroforests relative to natural forests. The results confirm that epiphyte diversity, composition, and vertical distributions are useful indicators of human disturbance and showed that while the studied plantations serve to preserve portions of epiphyte diversity in the landscape, their presence does not fully compensate for the loss of forests.     

Neither host-specific nor random: vascular epiphytes on three tree species in a Panamanian lowland forest

BACKGROUND AND AIMS: A possible role of host tree identity in the structuring of vascular epiphyte communities has attracted scientific attention for decades. Specifically, it has been suggested that each host tree species has a specific subset of the local species pool according to its own set of properties, e.g. physicochemical characteristics of the bark, tree architecture, or leaf phenology patterns. METHODS: A novel, quantitative approach to this question is presented, taking advantage of a complete census of the vascular epiphyte community in 0.4 ha of undisturbed lowland forest in Panama. For three locally common host-tree species (Socratea exorrhiza, Marila laxiflora, Perebea xanthochyma) null models were created of the expected epiphyte assemblages assuming that epiphyte colonization reflected random distribution of epiphytes in the forest. KEY RESULTS: In all three tree species, abundances of the majority of epiphyte species (69-81 %) were indistinguishable from random, while the remaining species were about equally over- or under-represented compared with their occurrence in the entire forest plot. Permutations based on the number of colonized trees (reflecting observed spatial patchiness) yielded similar results. Finally, a third analysis (canonical correspondence analysis) also confirmed host-specific differences in epiphyte assemblages. In spite of pronounced preferences of some epiphytes for particular host trees, no epiphyte species was restricted to a single host. CONCLUSIONS: The epiphytes on a given tree species are not simply a random sample of the local species pool, but there are no indications of host specificity either.     

Proximity is not a proxy for parentage in an animal-dispersed Neotropical canopy palm

We used parentage analysis to estimate seedling recruitment distances and genetic composition of seedling patches centred around reproductive trees of the animal-dispersed Neotropical canopy palm Iriartea deltoidea in two 0.5 ha plots within second-growth forest and one 0.5 ha plot in adjacent old-growth forest at La Selva Biological Field Station in north-eastern Costa Rica. Seedlings were significantly spatially aggregated in all plots, but this pattern was not due to dispersal limitation. More than 70 per cent of seedlings were dispersed at least 50 m from parent trees. Few seedlings were offspring of the closest reproductive trees. Seedling patches observed beneath reproductive trees originate from dozens of parental trees. Observed patterns of seedling distribution and spatial genetic structure are largely determined by the behaviour of vertebrate seed dispersers rather than by spatial proximity to parental trees.     

Retention of Inorganic Nitrogen by Epiphytic Bryophytes in a Tropical Montane Forest1

We developed and evaluated a model of the canopy of a tropical montane forest at Monteverde, Costa Rica, to estimate inorganic nitrogen (N) retention by epiphytes from atmospheric deposition. We first estimated net retention of inorganic N by samples of epiphytic bryophytes, epiphyte assemblages, vascular epiphyte foliage, and host tree foliage that we exposed to cloud water and precipitation solutions. Results were then scaled up to the ecosystem level using a multilayered model of the canopy derived from measurements of forest structure and epiphyte mass. The model was driven with hourly meteorological and event‐based atmospheric deposition data, and model predictions were evaluated against measurements of throughfall collected at the site. Model predictions were similar to field measurements for both event‐based and annual hydrologic and inorganic N fluxes in throughfall. Simulation of individual events indicated that epiphytic bryophytes and epiphyte assemblages retained 33–67 percent of the inorganic N deposited in cloud water and precipitation. On an annual basis, the model predicted that epiphytic components retained 3.4 kg N ha/yr, equivalent to 50 percent of the inorganic N in atmospheric deposition (6.8 kg N ha/yr). Our results indicate that epiphytic bryophytes play a major role in N retention and cycling in this canopy by transforming highly mobile inorganic N (ca. 50% of atmospheric deposition is NO^?₃) to less mobile (exchangeable NH⁺₄) and recalcitrant forms in biomass and remaining litter and humus.     

Diversity and Vertical Distribution Characteristics of Vascular Epiphytes in Bulong Nature Reserve Mengsong Section,Xishuangbanna

The first survey of vascular epiphytes was conducted using ground based inventory assisted by single rope technique in the recently established Bulong Nature Reserve, Xishuangbanna, China. Results indicated that vascular epiphytes were abundant and diverse there. On a total of 77 phorophytes in six plots (96 trees were examined in total, covered ca. 02ha area), 1756 individuals were recorded and were identified to 103 species (47 genera, 14 families). Compared with other regions, the epiphytes were as diverse as Paleotropics, and more diverse than temperate zone, but significantly less than the Neotropics. Orchids and ferns comprised 60%, 24% of the total flora, respectively, while others only took up 16%. The highest species richness and richest life form diversity was found in the middle canopy zone from 10 to 15m (51% of total species), where also supported high individual abundance (19% of total individuals). Besides the middle canopy, the most abundant zone of epiphyte individuals was detected at the base of the trunk (zone 0-5m, 24% of total individuals and 37% of total species), indicating another important niche for epiphytes in this forest environment. Primary hemiepiphytic figs were rare in the area and were not found on the surveyed host trees, while hemiepiphytic Araliaceae species (Schefflera elliptica and Tupidanthus calyptratus) were popular.     

Differences in epiphyte biomass and community composition along landscape and within-crown spatial scales

Vascular epiphytes contribute to the structural, compositional, and functional complexity of tropical montane cloud forests because of their high biomass, diversity, and ability to intercept and retain water and nutrients from atmospheric sources. However, human-caused climate change and forest-to-pasture conversion are rapidly altering tropical montane cloud forests. Epiphyte communities may be particularly vulnerable to these changes because of their dependence on direct atmospheric inputs and host trees for survival. In Monteverde, Costa Rica, we measured vascular epiphyte biomass, community composition, and richness at two spatial scales: (1) along an elevation gradient spanning premontane forests to montane cloud forests and (2) within trees along branches from inner to outer crown positions. We also compared epiphyte biomass and distribution at these scales between two different land-cover types, comparing trees in closed canopy forest to isolated trees in pastures. An ordination of epiphyte communities at the level of trees grouped forested sites above versus below the cloud base, and separated forest versus pasture trees. Species richness increased with increasing elevation and decreased from inner to outer branch positions. Although richness did not differ between land-cover types, there were significant differences in community composition. The variability in epiphyte community organization between the two spatial scales and between land-cover types underscores the potential complexity of epiphyte responses to climate and land-cover changes.     

The relationship between tree size and epiphyte species richness: testing four different hypotheses

Aim For epiphytic plants trees are habitat units, and tree size determines epiphyte species richness. While growing, trees generate vertical microhabitats that are exploited by epiphytes. One would expect to find four different types of relationship between tree size and epiphyte species richness: positive linear (young trees), neutral (old trees), negative (old decaying trees) and positive asymptotic (trees of mixed size class in a mature forest). We tested these relationships in plots of colonizing sweetgum trees in pastureland, isolated remnant trees in pastureland (old oaks) and sweetgum and oaks in a pristine forest. Location The study was carried out in a landscape shaped by the fragmentation of lower montane cloud forest in San Andrés Tlalnelhuayocan (19°30′56′′ N and 96°59′50′′ W; 1500–1600 m a.s.l.) in central Veracruz, Mexico. Methods We measured the d.b.h. of all oaks and sweetgum trees (d.b.h. ≥ 5 cm) present in pastureland and in three 100 m² plots of a lower montane cloud forest. All trees were climbed and species richness of the epiphytes recorded. Results As expected, colonizer trees in pastureland showed a linear positive relationship. Although we found evidence that remnant oaks in pastureland had a neutral relationship between tree size and epiphyte species richness, the low power of the test did not allow us to make conclusions about the kind of relationship. Mixed size‐class pristine forest trees showed a positive linear relationship between tree size and epiphyte species richness instead of a positive asymptotic one. Main conclusions Our results suggest that in the study area epiphyte communities are unsaturated, as the number of species increases with tree size and does not reach a ceiling. This evidence supports the idea that the species–area relationship is not asymptotic. However, the epiphyte community on remnant pastureland oaks may be saturated as epiphyte species richness did not increase with tree size, but a larger sample size is needed to confirm the neutral pattern. Neutral, asymptotic and negative patterns in the relationship between tree size and epiphyte species richness depend on the saturation of the trees by epiphytes. Other studies have suggested tree saturation, but further research is necessary in order to confirm or rule out these patterns.     

Rainforest air-conditioning: the moderating influence of epiphytes on the microclimate in tropical tree crowns

Epiphytes are often assumed to influence the microclimatic conditions of the tree crowns that they inhabit. In order to quantify this notion, we measured the parameters "temperature" (of the substrate surface and the boundary layer of air above it), "evaporative drying rate" and "evapotranspiration" at various locations within tree crowns with differing epiphyte assemblages. The host tree species was Annona glabra, which was either populated by one of three epiphyte species (Dimerandra emarginata, Tillandsia fasciculata, or Vriesea sanguinolenta) or was epiphyte-free. We found that during the hottest and driest time of day, microsites in the immediate proximity of epiphytes had significantly lower temperatures than epiphyte-bare locations within the same tree crown, even though the latter were also shaded by host tree foliage or branches. Moreover, water loss through evaporative drying at microsites adjacent to epiphytes was almost 20% lower than at exposed microsites. We also found that, over the course of several weeks, the evapotranspiration in tree crowns bearing epiphytes was significantly lower than in trees without epiphytes. Although the influence of epiphytes on temperature extremes and evaporation rates is relatively subtle, their mitigating effect could be of importance for small animals like arthropods inhabiting an environment as harsh and extreme as the tropical forest canopy.     

Effect of isolation on the structure and nutrient content of oak epiphyte communities

Because isolated trees in pasture experience greater exposure than forest trees, the epiphytes on them should be more drought and sun-tolerant. In Veracruz, Mexico, we compared the structure and nutrient content of the epiphyte community on five forest oaks (FO) in a fragment of lower montane cloud forest to that of five isolated oaks in pasture (IO). IO supported fewer epiphyte species than FO; 62.8% of the 35 epiphyte species were recorded only in one habitat (51.4% on FO and 11.4% on IO). Polypodium plebeium and Tillandsia spp. seedlings were more abundant on FO, while T. kirchhoffiana and T. punctulata were more abundant on IO. Evenness was lower on IO, which supported higher epiphyte biomass. pH, Ca, Mg, N, K, and Na concentrations were similar for FO and IO, with only P_extractable being lower on IO than FO. We concluded that when an epiphyte community is isolated (IO), the populations of some species expand while those of other diminish or disappear, a phenomenon that changes the structure of the epiphyte community becoming less even.     

Branchfall as a Demographic Filter for Epiphyte Communities: Lessons from Forest Floor-Based Sampling

Local variation in the abundance and richness of vascular epiphytes is often attributed to environmental characteristics such as substrate and microclimate. Less is known, however, about the impacts of tree and branch turnover on epiphyte communities. To address this issue, we surveyed branches and epiphytes found on the forest floor in 96 transects in two forests (Atlantic rainforest in Brazil and Caribbean rainforest in Panama). In the Brazilian forest, we additionally distinguished between edge and core study sites. We quantified branch abundance, epiphyte abundance, richness and proportion of adults to investigate the trends of these variables over branch diameter. Branches <2 cm in diameter comprised >90% of all branches on the forest floor. Abundance and richness of fallen epiphytes per transect were highest in the Brazilian core transects and lowest in the Panamanian transects. The majority of epiphytes on the floor (c. 65%) were found attached to branches. At all three study sites, branch abundance and branch diameter were negatively correlated, whereas epiphyte abundance and richness per branch, as well as the proportion of adults were positively correlated with branch diameter. The relationship between branch diameter and absolute epiphyte abundance or richness differed between study sites, which might be explained by differences in forest structure and dynamics. In the Panamanian forest, epiphytes had been previously inventoried, allowing an evaluation of our surveying method by comparing canopy and forest floor samplings. Individuals found on the forest floor corresponded to 13% of all individuals on branches <10 cm in diameter (including crowns), with abundance, richness and composition trends on forest floor reflecting canopy trends. We argue that forest floor surveys provide useful floristic and, most notably, demographic information particularly on epiphytes occurring on the thinnest branches, which are least accessible. Here, branchfall acts as an important demographic filter structuring epiphyte communities.