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.     

A First Quantitative Census of Vascular Epiphytes in Rain forests of Colombian Amazonia

Epiphytism in Colombian Amazonia was described by counting vascular epiphytes in thirty 0.025-ha (5 × 50 m) plots, well-distributed over the main landscape units in the middle Caquetá area of Colombian Amazonia. Each plot was directly adjacent to a 0.1-ha plot at which the species composition of trees and lianas (diameter at breast height (DBH) 2.5 cm) had been recorded 3 years earlier. The purpose of the study was to explore abundance, diversity, and distribution of epiphytes between the principal landscape units. A total of 6129 individual vascular epiphytes were recorded belonging to 27 families, 73 genera, and 213 species (which included 59 morpho-species). Araceae, Orchidaceae, and Bromeliaceae were the most speciose and abundant families. A total of 2763 phorophytes were registered, 1701 (62%) of which with DBH 2.5 cm. About 40–60% of the woody plants with DBH 2.5 cm carried epiphytes, which points at low phorophyte limitation throughout all landscapes. Epiphytism was concentrated on stem bases. Just as trees, epiphyte species assemblages were well associated with the main landscapes. Contrary to trees, however, epiphyte abundance and diversity (species richness, Fishers alpha index) hardly differed between the landscapes. This calls for caution when explanations for distribution and dynamics of tree species are extrapolated to growth forms with a totally different ecology.     

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.     

Biodiversity and spatial distribution of vascular epiphytes in two biotopes of the Cameroonian semi-deciduous rain forest

The diversity and spatial distribution of vascular epiphytes were surveyed in two biotopes (dryland forest and swamp-inundated forest) of the semi-deciduous rain forest area in Cameroon. Eight sites in each biotope were selected, which included 530 individuals of phorophytes in dryland forest and 460 in swamp-inundated forest. A total of 148 epiphyte species were recorded, which showed that semi-deciduous rain forests represent a major source of African-epiphyte diversity. Dryland forest hosted 110 epiphyte species, while the swamp-inundated forest harboured 108. A total of 56 species were found only in the dryland forest and 60 were restricted to the swamp-inundated forest. At family level, Orchidaceae exhibited the highest-species richness within both biotopes. Pteridophytes were more abundant in the swamp-inundated forest. A TWINSPAN analysis of the floristic similarities separated the two investigated forest types very clearly. No significant difference existed between the two biotopes regarding vertical distribution of epiphytes within single trees. The swamp-inundated forests may serve as a refuge for many epiphytes that occur in the outer canopy of the dryland forests, both because they are inherently less vulnerable to timber extraction, and enjoy better protection by legislation.     

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.     

Patterns in the diversity and distribution of epiphytes and vines in a New Zealand forest

Abstract Plants that rely on other plants for support (i.e. epiphytes and vines) are common in many forest ecosystems. However, they are poorly understood relative to terrestrial plants, especially in the Southern Hemisphere. To help bridge this gap, we evaluated the diversity and distribution of vascular epiphytes and vines on seven common tree species in a conifer‐broadleaf forest on New Zealand's North Island. Ground‐based surveys of 274 host trees were used to test whether epiphyte and vine diversity increased with tree diameter, and whether diversity‐diameter relationships differed among host tree species. Occurrence patterns of individual epiphyte and vine species were also assessed. We first evaluated the accuracy of ground‐based inventories by comparing surveys of trees made from the ground to those made from a canopy walkway. On average, 1 in 10 species of epiphytes and vines were unseen from the ground. However, sampling accuracy did not differ among the three host tree species growing along the walkway, suggesting unbiased comparisons could be made between hosts. Results from ground‐based surveys showed that species diversity of epiphytes and vines increased with host tree diameter. However, epiphytes showed stronger diversity‐diameter relationships than vines. Epiphyte diversity increased markedly in four host species and less strongly in the remaining three host species. Conversely, vines showed weak diversity‐diameter relationships in all host species. Occurrence patterns of individual species helped to explain diversity‐diameter relationships. All common epiphyte species occurred more frequently on large trees, regardless of host species, but occurrence patterns in most vine species were unrelated to tree size. Rather, the vines often showed strong host ‘preferences’. Overall results illustrate a rich diversity of distributional patterns in New Zealand's epiphytes and vines, and suggest that a similarly diverse set of ecological and evolutionary processes are responsible for them.     

Effect of Host Tree Traits on Epiphyte Diversity in Natural and Anthropogenic Habitats in Ecuador

Vascular epiphytes represent a highly diverse element of tropical rain forests, but they depend strongly on the structure and taxonomic composition of their tree communities. For conservation planning, it is therefore critical to understand the effect of host tree characteristics on epiphyte species richness in natural and anthropogenically transformed vegetation. Our study compares the effect of human land‐use on epiphyte diversity based on 220 study plots in a lowland rain forest and an Andean cloud forest in western Ecuador. We evaluate the relevance of host tree size and taxonomic identity for epiphyte species richness in contiguous primary forests, forest fragments, isolated remnant trees (IRTs), and secondary forests. At both study sites, epiphyte diversity was highest in primary forests, and it was lowest on IRTs and in secondary forests. Epiphyte species numbers of forest fragments were significantly reduced compared with the contiguous primary forest at the lowland study site, but not in the cloud forest area. Host tree size was a core predictor among secondary forests, but it had less significance within other habitat types. Taxonomic identity of the host trees also explained up to 61 percent of the variation in epiphyte diversity, especially for IRTs. The structural and taxonomic composition of the tree community in anthropogenically transformed habitat types proved to be fundamental to epiphyte diversity. This highlights the importance of deliberate selection of tree species for reforestation in conservation programs and the possible negative effects of selective logging in primary forests. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Epiphytes of Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis>) plantations in Ireland and the effects of open spaces

The epiphytes of the trunks and branches of mature Sitka spruce (Picea sitchensis) trees were studied in twelve plantations containing open spaces (glades, rides and roads) in the east and southwest of Ireland. A pair of trees was studied at each site: one tree at the south-facing edge of an open space and one in the forest interior. Spruce trees were found to support a moderately diverse range of bryophytes and lichens, including two relatively rare bryophyte species. Clear patterns in vertical distribution were identified, with bryophyte richness and cover decreasing and lichen richness and cover increasing from the tree base to the upper trunk. The open spaces themselves did not appear to affect overall epiphyte diversity, with no significant differences in any of the diversity measures between edge and interior trees. The main effect of open spaces was on the epiphyte cover of the edge trees. This was related to increased light levels combined with the presence of live branches from close to ground level on the south sides of the edge trees, which produced optimum conditions for bryophytes at the tree base and lichens in the upper plots. However, this dense side-canopy negatively affected epiphyte diversity on the north sides of the edge trees. Further research is required to assess the effects of open spaces within forestry plantations on epiphyte diversity.     

Epiphytes of Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis>) plantations in Ireland and the effects of open spaces

The epiphytes of the trunks and branches of mature Sitka spruce (Picea sitchensis) trees were studied in twelve plantations containing open spaces (glades, rides and roads) in the east and southwest of Ireland. A pair of trees was studied at each site: one tree at the south-facing edge of an open space and one in the forest interior. Spruce trees were found to support a moderately diverse range of bryophytes and lichens, including two relatively rare bryophyte species. Clear patterns in vertical distribution were identified, with bryophyte richness and cover decreasing and lichen richness and cover increasing from the tree base to the upper trunk. The open spaces themselves did not appear to affect overall epiphyte diversity, with no significant differences in any of the diversity measures between edge and interior trees. The main effect of open spaces was on the epiphyte cover of the edge trees. This was related to increased light levels combined with the presence of live branches from close to ground level on the south sides of the edge trees, which produced optimum conditions for bryophytes at the tree base and lichens in the upper plots. However, this dense side-canopy negatively affected epiphyte diversity on the north sides of the edge trees. Further research is required to assess the effects of open spaces within forestry plantations on epiphyte diversity.     

The epiphyte vegetation of Annona glabra on Barro Colorado Island, Panama

Aim Information on the community composition, structure, and dynamics of epiphyte vegetation is scarce. A survey of the epiphytes occurring on all individuals of one particular host tree species in a well-studied neotropical research site allowed us a comparison of the epiphyte flora of this tree with the local epiphyte flora, the analysis of spatial distribution patterns and the use of these patterns as indications for changes in time. In the future, our results can be used as a baseline data-set for the direct observation of the long-term dynamics in epiphyte communities. Location The study was conducted on Barro Colorado Island (BCI), Panama. Methods We recorded all individuals of the vascular epiphytes growing on Annona glabra L., a flood-tolerant, multiple-stemmed tree, which is restricted to the shoreline of BCI. Data on tree biometrics, epiphyte species, and epiphyte abundances were collected for more than 1200 trees. Results In total, we encountered almost 15,000 epiphytic individuals in sixty-eight species, corresponding to more than one third of the entire epiphyte flora of Barro Colorado Island. The component species differed strongly in abundance: the four most important species accounted for >75% of all individuals. In most cases, the same four species were also the first to colonize a tree (=phorophyte). Colonization patterns indicated no replacement of early colonizers by late arrivals. Species richness and epiphyte abundances showed a positive correlation with the size and the density of the host trees. All species showed a highly clumped distribution and the physiognomy of epiphyte communities of individual trees was dominated either by one or several of the four most common species or by a set of frequently co-occurring tank bromeliads. Other species were dominant only in exceptional cases. Most species were always rare. A distance effect on community composition was mostly confined to a local scale with an increased similarity in the species assemblage of stems of a tree v. neighbouring trees. Main conclusions The epiphytes on a single small phorophyte species may encompass a surprisingly large proportion of the local epiphyte flora. The observations that most tree crowns are inhabited by a single or only very few species, and that all epiphyte species show highly clumped distributions suggest a predominance of very local dispersal within a tree crown, which is only infrequently interrupted by successful long-distance dispersal between crowns.     

Differential diameter-size effects of forest management on tree species richness and community structure: implications for conservation

In this paper we tested the hypothesis that logging effects in the adult tree community reverberate upon the regeneration contingent. We examined the differences on the tree community between forest reserves and 10 year-old logged areas in the Yucatan Peninsula. We used a paired design in three independent sites to estimate the effects of logging on tree species richness, diversity, composition and structure. Analyses were conducted differentiating individuals of four diameter-size classes: 1–5, 5–10, 10–25, and >25 cm DBH. We found out that there were differential effects by size. Species richness in the smaller and larger diameter-size classes was significantly lower in logged areas. Floristic composition was also different between logged and unlogged areas, with a trend towards more secondary forest associated species and less primary forest associated species in logged areas, and a higher density of species represented by a single individual in unlogged reserves. In terms of structure, trees DBH <10 cm, lianas, and re-sprouting stumps were more abundant in logged areas. Our findings suggest that 10 years after logging, harvested areas show alterations in structure, and potentially a reduction in species richness. We suggest that to make timber extraction and forest conservation compatible at this site, it is necessary to gain a better understanding of the ecology and regeneration requirements of the less abundant species, and to assess whether current logging practices might hinder their permanence in the study area.     

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.     

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。     

Diversity and biogeography of vascular epiphytes in Western Amazonia, Yasuní, Ecuador

Aim Although vascular epiphytes are important components of species richness and complexity of Neotropical forests, vascular epiphytes are under‐represented in large scale biogeographical analyses. We studied the diversity, biogeography and floristic relationships of the epiphytic flora of the Yasuní region (Western Amazonia) in a Neotropical context, with special emphasis on the influence of the Andean flora on floristic composition and diversity of surrounding lowland forests. Location Western Amazonian lowland rainforest, Tiputini Biodiversity Station (0°38′ S 76°09′ W, 230 m a.s.l., 650 ha), Yasuní National Park, Ecuador. Methods We compared the vascular epiphyte flora of Yasuní with 16 published Neotropical epiphyte inventories. Secondly, based on a floristic database with records of more than 70,000 specimens of vascular epiphytes from the Neotropics the elevational composition of eight selected inventories was analysed in detail. Results The vascular epiphyte flora of Yasuní is characterized by a very high species richness (313 spp.). A moderate portion of species is endemic to the Upper Napo region (c. 10%). However, this figure is much higher than previous analyses primarily based on woody species suggested. Geographical ranges of these species match with a proposed Pleistocene forest refuge. Compared with Northern and Central Amazonian sites, Western Amazonian epiphyte communities are characterized by a higher portion of montane and submontane species. Species richness of vascular epiphytes at the sites was correlated with the amount of rainfall, which is negatively correlated with the number of dry months. Main conclusion Recent and historic patterns of rainfall are the driving forces behind diversity and floristic composition of vascular epiphytes in Western Amazonia: high annual rainfall in combination with low seasonality provides suitable conditions to harbour high species richness. The proximity to the Andes, the most important centre of speciation for most Neotropical epiphytic taxa, in combination with the climatic setting has allowed a continuous supply of species richness to the region. At least for epiphytes, the borderline between the Andean and Amazonian flora is much hazier than previously thought. Moreover, the comparatively moist climate in Western Amazonia during the Pleistocene has probably led to fewer extinctions and/or more speciation than in more affected surrounding lowlands.     

Diversity and distribution of herbaceous vascular epiphytes in a tropical evergreen forest at Varagalaiar, Western Ghats, India

Herbaceous vascular epiphytes were screened in a total sample of 13 445 trees (in 153 species) and 348 lianas (in 30 species) 30 cm girth at breast height in a 30 ha plot of tropical evergreen forest at Varagalaiar, Indian Western Ghats. Of these, 4.3% of trees and 3.7% of lianas were infested with epiphytes. Epiphyte diversity totaled 26 species in 19 genera and 10 families. Sixteen species were angiosperms in three families (Orchidaceae 54%; Piperaceae and Araceae 8%) and 10 species (38%) were pteridophytes in seven families. The orchid, Pholidota pallida was most abundant and occurred on 178 (29.6%) stems. Asplenium nidus occurred on the maximum of 62 host species. The species richness estimators employed for species accumulation curves after 100 times randomization of sample order have stabilized the curve at 18th and 19th hectares respectively for Incidence-based Coverage Estimator and Chao2. A total of 588 trees and 13 liana stems lodged epiphytes, 74% of which were evergreen species and 26% deciduous. Epiphyte density was greater (56%) in deciduous species (Bischofia javanica 30% and Vitex altissima 8%). A significant positive relationship was found between trunk size and epiphyte association. Larger epiphyte species occurred mostly on middle and larger stems and smaller epiphyte species occurred on smaller stems. The majority of epiphytes (92%) were of autochorous dispersal type, bearing capsule or dust diaspores, while the remaining species with berries and nutlets are dispersed by small vertebrates.     

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.     

Vascular Epiphytes on Isolated Pasture Trees Along a Rainfall Gradient in the Lowlands of Panama

Secondary habitats are increasing in importance in tropical countries due to ongoing destruction of pristine vegetation. In spite of the magnitude of current changes, our understanding of their effects on nontrees (e.g., nonvascular or vascular epiphytes) is still very patchy, particularly in lowland habitats. Here, we report a study with isolated pasture trees in southwest Panama. The >800 studied trees, which belonged to >100 different species, harbored almost 27,000 epiphytes of 83 species. Orchidaceae was the most species‐rich family, with almost 60 percent of all species, while Bromeliaceae were most abundant. A rainfall gradient in the study region from ca 1000 to >3000 mm explained more of the variation in species abundance and richness than host characteristics (e.g., species identity, tree size). The unexpectedly large number of epiphytes in these pastures still represents a substantial change relative to a natural setting, which is suggested by a comparison with a forest inventory under similar climatic conditions. In pastures, species richness was lower as deduced from individual‐based rarefaction curves, a larger proportion of species and individuals showed crassulacean acid metabolism, and the relationship of epiphyte abundance/species richness and tree diameter was much less steep. Even the already reduced diversity, however, may be only transient in secondary habitats—the long‐term persistence of epiphyte populations in pastures is an open question and has to be addressed by repeated monitoring to fully evaluate the significance of pasture trees for the conservation of vascular epiphytes in tropical lowlands.     

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.     

Contrasting functional traits maintain lichen epiphyte diversity in response to climate and autogenic succession

Aim Lichen epiphytes are important for biodiversity conservation and are also widely applied as environmental indicators. However, biogeographical and ecological knowledge underpinning lichen epiphyte conservation, and the use of lichens as indicators, is based primarily on a limited range of ‘macrolichen’ species. Wider trends in epiphyte biodiversity remain largely unexplored. This paper examines the community structure of lichen epiphytes on aspen (Populus tremula L.) in Scotland, including species across all functional groups and comprising, therefore, taxonomically difficult ‘microlichens’. Location Northern Britain (Scotland). Methods Epiphytes were sampled from 12 sites throughout Scotland and examined at two scales: between and within aspen stands. Species were classified into contrasting functional groups and ordination by detrended correspondence analysis was used to summarize community structure. Results Within aspen stands (between trees) epiphyte communities showed successional patterns related to tree age. These successional patterns changed predictably for stands aligned along a climatic gradient (between stands). Main conclusions A dual climatic–successional trend in epiphyte community structure is presented. Large‐scale trends in epiphyte diversity are explained as the local response of species with contrasting functional traits to climate and autogenic succession. Turnover of functional groups between stands is positively related to β‐diversity, and ecological limits to the frequency of contrasting functional groups are presented. Accordingly, the study and application of lichen species with similar functional traits may inadequately represent patterns of biodiversity. This prompts criticism of the currently accepted conservation strategy, i.e. (1) an emphasis in the conservation literature on ‘macrolichen’ species with similar ecologies and (2) the application of lichen indices over climatically variable geographical areas.     

Primary ecological succession in vascular epiphytes: The species accumulation model

Epiphytes are integral to tropical forests yet little is understood about how succession proceeds in these communities. As trees increase in size they create microhabitats for late‐colonizing species in both small and large branches while maintaining small tree microhabitats for early colonizing species in the small and young branches. Thus, epiphyte succession may follow different models depending on the scale: at the scale of the entire tree, epiphytes may follow a species accumulation model where species are continuously added to the tree as trees increase in size but at the scale of one zone on a branch (e.g., inner crown: 0–2 m from the trunk), they may follow the replacement model of succession seen in terrestrial ecosystems. Assuming tree size as an indicator of tree age, I surveyed 61 Virola koschnyi trees of varying size (2.5–103.3 cm diameter at breast height) in lowland wet tropical forest in Costa Rica to examine how epiphyte communities change through succession. Epiphyte communities in small trees were nested subsets of those in large trees and epiphyte communities became more similar to the largest trees as trees increased in size. Furthermore, epiphyte species in small trees were replaced by mid‐ and late‐successional species in the oldest parts of the tree crown but dispersed toward the younger branches as trees increased in size. Thus, epiphyte succession followed a replacement model in particular zones within treecrowns but a species accumulation model at the scale of the entire tree crown.