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 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.     

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.     

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。     

Vertical stratification of vascular epiphytes in submontane and montane forest of the Bolivian Andes: the importance of the understory

We studied species richness, composition, and vertical distribution of vascular epiphytes at two sites in the Bolivian Andes. To account for the epiphyte flora on understory trees, epiphytes on shrubs and small trees were sampled in 20 × 20 m² subplots around each sampled canopy tree; this understory zone U is introduced as an addition to the well-established five vertical Johansson tree zones. More than 20% of about 500 species recorded were found only in the understory subplots, including ca. 40% of aroids, 35%–40% of piperoids, and 25%–30% of ferns. Habitat generalists (occurring in three or more zones) were most common, contributing about 50% of all species, specialists (occurring only in two zones, or in three continuous ones) 34%–42%, and hemiepiphytes 6%–16%. Canopy epiphytes (occurring > 90% in tree zones Z3–5) were mainly represented by orchids and ferns, many with special adaptations to drought stress such as pseudobulbs, succulence, and poikilohydry. Trunk epiphytes ( > 90% in understory and tree zones Z1–2) reached highest relative species numbers among piperoids and ferns. Most hemiepiphytes were also trunk epiphytes, due to their characteristic growth pattern, and included mainly aroids. The vertical distribution of epiphytes within a tree is determined by several microenvironmental gradients, with light intensity, wind speed, and air temperature increasing and air humidity decreasing from the ground level to the canopy.     

Sponge–seaweed associations in species of Ptilophora (Gelidiaceae, Rhodophyta)

Sponge–seaweed associations in the seaweed genus Ptilophora are poorly understood; therefore, 94 specimens, representing all 17 species of Ptilophora, were examined to detail this phenomenon. All but 2 Ptilophora species were shown to produce surface proliferations, with 13 species found to have sponge associations. Evidence for facultative sponge epiphytism was found with species–specific interactions being unlikely. Results show that surface proliferations are not induced by sponge epiphytes, as they often occur in the absence of sponge epiphytes, and vice versa. The significant number of proliferate thalli found with sponge epiphytes suggests that there is a likely relationship between the presence of surface proliferations and sponge infestation. Sponge epiphytes and Ptilophora species appeared structurally related in that the sponge probably exploits a niche habitat provided by the alga, for which surface proliferations might aid the sponge in bonding to the alga.     

Vascular Epiphyte Communities in the Inner-Crown of Hyeronima alchorneoides and Lecythis ampla at La Selva Biological Station, Costa Rica

The community composition and percent cover of vascular epiphytes were examined in relation to photosynthetic photon flux density (PFD), temperature, vapor pressure, and tree characteristics in the inner-crown of two emergent tree species, Hyeronima alchorneoides and Lecythis ampla, at La Selva Biological Station, Costa Rica. A total of 53 species were found in a total sampled branch area of 32 m² in eight trees—four trees per species. Community composition varied both among individuals of the same tree species and between tree species. However, percent cover patterns of vascular epiphytes were significantly different between the two tree species; Hyeronima had a significantly greater percent cover of epiphytes than Lecythis . The higher percent light transmittance as well as lower humidity in Lecythis are likely causes of its lower percent cover of epiphytes.     

The floristic compositions of vascular epiphytes of a seasonally inundated forest on the coastal plain of Ilha do Mel Island, Brazil

A 3,000 m2 area of seasonally inundated forest on the island of Ilha do Mel (25 degrees 30" S 48 degrees 23" W) in Paraná, Brazil, was sampled by collecting plants from all strata, using climbing equipment when necessary. The area harbors 103 species of epiphytes, in 49 genera and 20 families, of which 28 species are pteridophytes and 75 magnoliophytes (64 Liliopsida, 11 Magnoliopsida). The most common families are Orchidaceae, Bromeliaceae, Polypodiaceae and Araceae, and frequent genera are Vriesea, Epidendrum, Maxillaria, Pleurothallis and Prosthechea. Eight families were represented by one species each. Most species were classified as obligatory holoepiphytes (62 %), followed by the relatively more rare preferential holoepiphytes (13 %), facultative epiphytes (11 %), hemiepiphytes (9 %) and accidental epiphytes (6 %).     

Species richness,composition, and spatial distribution of vascular epiphytes in Amazonian black-water floodplain forests

This study examines the occurrence of vascular epiphytic species in Central Amazonian black-water floodplain forests (igapó) and considers whether their horizontal and vertical distribution is influenced by the flood pulse, as is the case with tree species (phorophytes). Research was conducted in sixteen forest plots the Jaú National Park. In these, epiphytes on all phorophytes with DBH ≥ 10 cm were identified. We measured flood height using the watermark left by the last high-water period, then estimated the height relative to the ground of every epiphytic individual. We recorded 653 individuals in 37 species, distributed on 109 phorophytes. Igapó floodplain forests have much lower richness and abundance of vascular epiphyte species than do other Amazonian forests. This may reflect the limitation of available sites for colonization (only 24.9% of studied trees were occupied by epiphytes). Holoepiphytes predominated, and the combined presence of a flood-pulse, linked to the nutrient-poor soil poor seems to limit the occurrence of nomadic vines. Horizontal distribution of epiphytes followed the distribution of phorophytes, which in turn followed the flood-level gradient. Also flooding interacted strongly with vertical zonation to determine species richness. As already well-reported for trees, and unlike reports of epiphytes in other floodplains, flooding strongly influenced richness and distribution of vascular epiphytes in the studied igapó forests.     

Effects of Multiple Disturbances (Fire and Hurricane) on Epiphyte Community Dynamics in a Subtropical Forest,Florida, U.S.A.1

We addressed the interacting effects of a natural large–scale fire and a subsequent major hurricane on relative positions of epiphytes in a subtropical forest. In Everglades National Park, subtropical hammocks (hardwood tree “islands”; burned and unburned) during the Ingraham Fire (1989) were surveyed for trees and epiphytic bromeliads (Tillandsia spp.) one year before, as well as one and five years after, Hurricane Andrew (1992). We measured trees (species, diameter, and status [alive/dead]) and epiphytes (species, height, host tree characteristics, substrate life status, and density). The fire decreased the height of epiphytes during the hurricane because branches and bark of trees killed by the fire were unstable epiphyte substrates in the high winds. Proportions of epiphytes on Quercus virginiana were equally increased after the hurricane in both unburned and burned hammocks; the large size and bark characteristics resulted in greater proportional survival of epiphytes on this species. During the five years following the hurricane, changes in the distributions of epiphytes generally were toward pre–hurricane distributions, but recovery was faster in unburned than burned hammocks. We conclude that disturbances that kill trees are likely to amplify the vertical reduction of epiphytes during a subsequent hurricane and that effects of a single disturbance on plant populations can be influenced by the disturbance history of the system, including different types of disturbances.     

Pteridophyte species richness in Andean forests in Bolivia

I assessed the magnitude and distribution of pteridophyte species richness on the eastern Andean slope in Bolivia based on 676 study plots of 400 m² each in forest habitats at 65 study sites. In total 755 species were recorded, including 426 (56%) epiphytes and 598 (79%) terrestrials, with 266 species (35%) recorded under both groups. Mean number of species per plot at a given site varied from 0 to 31.1 for epiphytes, 0 to 20.9 for terrestrials, and 0 to 47.9 for all species combined. The highest numbers of species recorded at a given study site were 110 epiphytes, 101 terrestrials, and 167 species in total. While overall there were more terrestrial than epiphytic species, at individual sites and plots the reverse was true, indicating that terrestrial species tended to be more patchily distributed than epiphytes. Despite high survey intensity, many species went unrecorded; the minimum estimate of total species richness obtained through extrapolation was 975 species overall, including 559 epiphytes and 880 terrestrials. A correlation analysis of species richness to 14 environmental parameters revealed a highly positive correlation to mean annual precipitation and bryophyte cover on tree branches (a proxy for air humidity). Significant correlations to other parameters (e.g. human impact, canopy height, etc.) reflected the covariance of these factors with precipitation and bryophyte cover. Despite a lack of data on the pteridophyte communities from much of the Bolivian Andes, it appears that in most of the countries, pteridophyte diversity can be protected by focussing the most humid parts of the Andean forests.     

Contribution of peeling host for epiphyte abundance in two tropical dry forests in the “El Cielo Biosphere Reserve”, Mexico

Host traits partly determine the abundance and species richness of epiphytes in tropical forests. It has been proposed that older trees with rough bark and evergreens often house more individuals and more epiphytic species than those with thin, smooth, and peeling bark, which harbor few epiphytes. We hypothesize (i) that epiphytes are more abundant and species-rich in the more shaded forest, which is related to bark roughness, and (ii) that epiphytes are distributed in the middle of the host, where microenvironmental conditions are more favorable to survival. We evaluated abundance, species richness, and vertical distribution of epiphytes in two tropical dry forests, according to the deciduousness and basal area of the trees. Moreover, we selected the most abundant epiphytes to test whether their distribution is related to a specific bark type and examine their vertical distribution in two dry forests. We distinguished a high abundance and species richness of epiphytes in the deciduous forest, although basal area and host species richness were higher in the semi-deciduous forest. In both forests, we found a positive relationship between epiphyte abundance and basal area. Higher abundance of epiphytes was related to the predominance of Tillandsia schiedeana, a drought-adapted species, in both forests. Unexpectedly, epiphytes abundantly colonized Bursera simaruba, a host with peeling bark and a very branched crown, where small individuals of T. schiedeana colonized abundantly toward the top of the crown. Our results show the importance of the tropical dry forest, particularly, B. simaruba, in maintaining epiphyte diversity in terms of T. schiedeana colonization.     

西双版纳热带森林附生维管植物的物种多样性与区系特征

附生植物是山地森林生态系统中重要的结构性成分。因受林冠调查技术限制, 人们对林冠附生植物知之甚少。本文在前人有关西双版纳植物区系研究的基础上, 结合野外调查和标本资料, 对该地区附生植物的物种组成与分布进行了整理。结果表明, 西双版纳热带森林附生维管植物共有29科134属486种, 约占全部维管植物的11%。附生兰科是最丰富的类群(69属293种), 其中以石斛属(Dendrobium) (49种)和石豆兰属(Bulbophyllum) (48种)物种数最多。蕨类是仅次于兰科的附生类群(13科38属97种), 其中水龙骨科(51种)占附生蕨类总数的50%以上。基于生活史和养分来源的划分标准, 该地区48%的附生物种属于兼性附生, 其次是以兰科为主的专性附生(46%), 半附生仅占6%。从区系上看, 西双版纳附生植物属的分布具有明显的热带亚洲性质。附生植物主要分布于资源受限的林冠生境, 对环境变化极为敏感, 在人为干扰日益加剧的背景下, 这类植物正面临严重威胁。因此, 需要加强对西双版纳林冠亚系统的保护。     

Pollination and phenology of flowers in the canopy of two contrasting rain forest types in Amazonia,Colombia

The main objective of this investigation was to study the pollination characteristics of two types of Amazonian rain forest at plant community level. Seasonally inundated forest was compared with upland (tierra firme) forest. The study focused on plant species in the canopy. The pollination spectra show that in both forests most canopy trees and lianas are pollinated by small bees, large bees, butterflies or by small, relatively unspecialized insects. In the upland forest small bees are the most important pollinators (32% of all species of trees and lianas are pollinated by them), whereas large bees are predominant in the floodplain (22%). Other pollinators, like hummingbirds, bats, moths, and beetles are less common (>10%), but always somewhat more important in the flood plain than in the upland forest. Bees are the most common pollinators of epiphytes. In the flood plain forest, flies are also important as epiphyte pollinators (19%), whereas in the upland forest hummingbirds pollinate more epiphytes. The phenological patterns are quite similar in both the upland and the flood plain. We found a peak in flowering in the transition period between the wet and the dry season. Flowering activity was lowest during the wet season. Differentiation in sexual systems was correlated with life form. Dioecy and monoecy were found mostly among tree species. Most species of all life forms though were hermaphroditic. No difference with respect to the relative importance of sexual systems was found between the two forest types.     

Johansson revisited: the spatial structure of epiphyte assemblages

Question: Vertical zonation schemes are widely used in biodiversity studies with vascular epiphytes as a tool to capture spatial distribution patterns, the one most commonly used was proposed by Johansson more than 30 years ago. Does a survey of the epiphytes found on larger trees really yield a representative sample of the local community? Location: Lowland rainforest of the San Lorenzo Crane Plot, Republic of Panama. Methods: A complete census of the vascular epiphytes on all trees > 1 cm DBH in 0.4 ha of undisturbed lowland forest was analysed with both cluster and discriminant analysis to detect groupings of epiphyte species. Results: Six different groups of species were detected, five of them preferring different substrates on larger trees (as defined by (1) the height above ground at the attachment site, (2) the diameter of the substrate and (3) the occurrence on stem vs branches/twigs) and resembling to some extent the original Johansson zones. A sixth group of epiphytes, comprising ca. 10% of all taxa, was almost always found on small diameter stems and branches of trees with small DBH at lower and intermediate heights within the forest. Conclusions: Applying pre‐established zonation schemes may lead to misleading results in biodiversity studies with epiphytes. Important aspects of spatial distribution patterns may be missed, and the determination of relative species abundances may carry a strong quantitative and qualitative bias when analyses rely completely on epiphytic plants found on larger trees.     

How prevalent is crassulacean acid metabolism among vascular epiphytes?

The occurrence of crassulacean acid metabolism (CAM) in the epiphyte community of a lowland forest of the Atlantic slope of Panama was investigated. I hypothesized that CAM is mostly found in orchids, of which many species are relatively small and/or rare. Thus, the relative proportion of species with CAM should not be a good indicator for the prevalence of this photosynthetic pathway in a community when expressed on an individual or a biomass basis. In 0.4 ha of forest, 103 species of vascular epiphytes with 13,099 individuals were found. As judged from the C isotope ratios and the absence of Kranz anatomy, CAM was detected in 20 species (19.4% of the total), which were members of the families Orchidaceae, Bromeliaceae, and Cactaceae. As predicted, the contribution of CAM epiphytes to the total number of individuals and to total biomass (69.6 kg ha^-1) was considerably lower (3.6% or 466 individuals and, respectively, 3.0% or 2.1 kg ha^-1).     

Diversity and phytogeography of vascular epiphytes in a tropical–subtropical transition island,Taiwan

We present the first checklist of vascular epiphytes in Taiwan, based on herbarium specimens, literature records, and field observations. Epiphyte phytogeography was analyzed using Takhtajan's modified division in floristic regions. We ascertain the presence of 336 species of vascular epiphytes (24 families, 105 genera) in Taiwan. Pteridophytes contribute most species (171 species), followed by orchids (120 species). Epiphytes contribute 8% to Taiwanese floristic diversity and epiphyte endemism is near 21.3%. The extensive mountain system is probably the most effective driver for epiphyte diversification and endemicity in Taiwan. Phytogeographically, Taiwanese epiphytes exhibit equal affinity to the Malesian region, southern China and Indo-China and Eastern Asiatic regions. However, some species have a disjunctive distribution between Taiwan and SW China and/or E Himalaya, presumably related to low habitat similarity with adjacent China and/or the legacy of Late Quaternary climate change. Vascular epiphyte distribution patterns corroborate the phytogeographical separation of the island of Lanyu from the main island of Taiwan along Kanto's Neo-Wallace Line.     

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.     

Do host bark traits influence trunk epiphyte communities?

Host bark traits are known to affect the characteristics of epiphyte communities in forests worldwide; however, few investigations of such relationships have been undertaken in New Zealand forests. By examining the trunk epiphyte communities on four co-occurring forest tree species (Agathis australis, Dacrydium cupressinum, Knightia excelsa and Vitex lucens) representing a range of bark characteristics, we sought evidence that bark traits may shape these communities. Sampling was conducted on tree trunks in the Waitakere and Hunua ranges in the Auckland Region. As expected, the rough but lightly shedding bark of Vitex lucens was found to support many epiphytes, whereas the coarsely flaking bark surface of Dacrydium cupressinum supported few epiphytes. Conversely, despite bark with a smooth texture that sheds in large flakes, and contrary to the suggestions of previous authors, Agathis australis trunks were found to support the greatest numbers of epiphytes and this species was one of the most frequent epiphyte hosts. The individual epiphytes found on Agathis australis, however, were significantly smaller and more appressed to the trunk than those on the other trees, and species composition differed from the other host species.