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。     

Vascular plant species count in a wet forest in the Chocó area on the Pacific coast of Colombia

The total number of vascular plant species was counted and growth form distribution was studied in the Chocó area on the Pacific coast of Colombia, in two transects 400×10m and ten transects 2×50m, for a total sampled area of 0.9ha. The species count of the ten transects (442 species in 0.1ha) appears to be the highest number of species recorded with this methodology. There were 970 species for the total area (0.9ha). Ninety to ninety-five percent of the species were under 10cm dbh and 70–86% under 2.5cm dbh, epiphytes and small trees and treelets 10cm dbh being the most diverse growth forms. The most species-rich families and genera were those represented by herbaceous plants and treelets. Individuals were counted only in the ten 2 × 50 m transects (0.1 ha), where 4459 individuals were found. Palms and ferns were the most abundant growth forms. Arguments are presented against the way diversity is usually measured. Recommendations are made to include other growth forms besides trees when assessing alpha diversity for conservation purposes.     

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.     

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。     

The vascular epiphytes of a lowland forest in Panama—species composition and spatial structure

We conducted a comprehensive census of the vascular epiphytes in a lowland forest in Panama by means of a canopy crane. In 0.4 ha of ca. 40-m tall forest, 103 species of vascular epiphytes with 13,099 individuals were found. The orchids were the most important family both in species and individual numbers, accounting alone for >40% of all species and >50% of all individuals. There was a clear vertical segregation of species with a concentration at intermediate heights: more than 50% of all individuals were found between 15 m and 25 m above ground. Tree species identity, tree size and the position of a tree in the forest (“space”) all influenced species composition. However, none of the two environmental variables nor space alone explained more than 10% of the total variation in epiphyte assemblages in several canonical correspondence analyses. By far the largest proportion of the observed variation remained unexplained and is arguably due to mere chance. In the future, our results will be used as a baseline data-set for the direct observation of the long-term dynamics in a diverse epiphyte community.     

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.     

亚热带森林附生植物叶片气孔特征及其可塑性对光照变化的响应

附生植物是热带亚热带森林生态系统中物种多样性极高且极其脆弱敏感的生物类群之一。光照被认为是促进附生植物由陆生类群演化而来并决定其生长和分布的关键因素。然而,由于接近林冠和规范性采样的限制,附生植物与光照的关系仍亟待阐述。为揭示附生植物对光强变化的响应和适应策略,该研究以亚热带常绿阔叶林6种附生植物(林冠层木本:鼠李叶花楸、毛棉杜鹃;林冠层草本:狭瓣贝母兰、毛唇独蒜兰;树干区草本:点花黄精、距药姜)为对象,对其在4个光处理梯度下生长的叶片气孔特征及其可塑性进行了对比分析。结果表明:(1) 2种附生小乔木的气孔面积(SA)、气孔密度(SD)、潜在气孔导度指数(PCI)和表皮细胞密度(ECD)均对光强改变显著响应。2种附生兰科植物的SA最大,而SD最小;附生乔木叶片SD和ECD的光响应趋势与陆生植物更相似,而附生草本则出现种间差异。(2) 6种附生植物的气孔、表皮细胞特性及其表型可塑性,在草本-木本、常绿-落叶植物、林冠-树干区之间,均无明显差别。(3)附生植物气孔特性和表皮细胞平均可塑性指数均低于陆生植物。综上结果表明,亚热带常绿阔叶林中附生植物对于光环境变化的适应性相对较弱。不同的附生植物可以通过不同程度地增加叶片SD和ECD来适应高光强生境,并通过对SD和SA的双重调节以增大潜在光合能力从而应对低光胁迫。     

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.     

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.     

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.     

A metapopulation approach to the analysis of long‐term changes in the epiphyte vegetation on the host tree Annona glabra

Question: Do vascular epiphyte species have a metapopulation structure? What are the qualitative and quantitative long‐term changes of the complete vascular epiphyte vegetation in a particular host tree species? Location: Lowland forest on Barro Colorado Island (9° 10’ N, 79°51’ W), Republic of Panama. Methods: In 1994 and 2002 we conducted a census of all vascular epiphytes growing on more than 1000 Annona glabra trees (= patches). Epiphyte species abundances were recorded at the tree level in each census. Results: The number of epiphyte individuals increased from ca. 15 000 to ca. 23 700 individuals during the census interval while the species composition on Annona glabra as a whole was rather stable. There was a strong positive relationship between occurrence in patches and local abundance of the species, and between species richness and host tree stand size. The dynamics of local populations of a given species were uncorrelated to each other; small and large local populations of most species had the same probability to go extinct. The frequency distribution of species on all host trees was not bimodal, but on a subset of heavily colonized host tree stands it was. Numbers of species and individuals were correlated with tree size which was not due to a correlation of tree size and tree age. Conclusions: As far as the most abundant epiphyte species with metapopulation structures are concerned, these species belong to diverse families, e. g. Orchidaceae, Bromeliaceae and Polypodiaceae. Even ca. 80 years after the initial establishment of the host tree species in the study area epiphytes are still in the stage of initial colonization and have not reached a steady state as indicated by the strong increase in individuals and the ongoing colonization of empty trees.     

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.     

The vertical distribution and abundance of gastropods and bivalves from rocky beaches of Cuastecomate Bay, Jalisco, México

The vertical distribution and abundance of conspicuous gastropod and bivalve species were studied at five rocky beaches in Cuastecomate Bay, Jalisco. Sampling was done from September, 1993 through March, 1994 with 0.75 m2 quadrants placed along replicate transect lines (10 m long) in the supralittoral and mesolittoral (upper, middle and lower intertidal) zones. A total of 6,643 mollusks were collected. Gastropods dominated the samples (6,272 individuals, 44 species); the bivalves were less abundant and diverse (371 individuals, five species). Seventeen species comprised 89.8% of all individuals collected. The gastropods Nodilittorina aspera and Nerita scabricosta were the most abundant with 637.8 and 71.43 individuals/m2, respectively. The most abundant bivalves were Brachidontes adamsianus and Chloromytilus palliopunctatus with 60.7 and 61.3 individuals/m2 respectively. The abundance of gastropods decreased from the supralittoral to the lower tidal zones while the number of species increased in the same direction. The number of species of bivalves also increased from the supralittoral to the lower intertidal zone; the abundance of individuals was higher at the middle intertidal zone. Affinities between groups of species among sampling stations were identified by computing Pearson's correlation coefficient using abundance values (ind./m2) and Jaccard's dissimilarity index using species presence or absence in the lower intertidal zone. Affinity among stations was not dependent upon their vicinity but on the high dominance of few species, the occurrence of many secondary species and beach characteristics.     

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.     

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.     

Effects of tree crown structure on biomass of the epiphytic fern Polypodium scouleri (Polypodiaceae) in redwood forests

Redwood forests contain some of the largest and most structurally complex trees on Earth. The most abundant vascular epiphyte in these forests is the fern Polypodium scouleri (Polypodiaceae). We measured dimensions of all 765 P. scouleri mats on 32 trees (27 Sequoia sempervirens, 5 Picea sitchensis). Eighteen P. scouleri mats from 11 trees were randomly selected for removal and dissection in the laboratory. The total fern mat mass consisted of live fronds (3.3%), dead fronds (2.4%), live rhizomes (4.2%), dead rhizomes (8.9%), roots (34.4%), humus (28.0%), and debris (18.8%). We used multiple regression analysis to develop equations for estimation of fern masses, and we applied these equations to undisturbed fern mats on the 32 trees. Individual trees supported up to 742 kg dry mass of P. scouleri mats. These are the highest whole-tree epiphyte masses ever reported. We also quantified crown structure and counted the number of vascular plant species occurring as epiphytes on each tree. Very large, complex trees had more fern mat mass and higher vascular epiphyte species richness than smaller, simpler trees. Desiccation-sensitive organisms dependent on water stored in fern mats may be unable to survive in managed redwood forests lacking large, complex trees with abundant P. scouleri.     

林冠附生植物生态学研究进展

林冠附生植物及其枯死存留物是构成山地湿性森林生态系统中生物区系、结构和功能的重要组分。由于在林冠攀爬技术上的限制,过去对林冠附生植物在生态系统结构和功能过程中的作用未能引起足够的重视。近20年来,随着对林冠生物多样性及其在生态系统功能过程影响的认识和研究技术上的提高,对林冠附生生物的研究已逐步从个体水平转移到系统水平上。有关林冠附生植物多样性、生物量及其生态学效应已成为近年来国际上新兴研究领域——“林冠学”的研究热点之一。许多研究表明,林冠附生植物在生态系统水平上的交互作用比它们的解剖、形态和生理特征更为重要。国外大量的研究结果表明,林冠是一个适合于许多生物种类生存的场所,其数量比想象的更为丰富。在全球范围内估计有29 500余种附生植物,其中维管束附生植物的种类高达24 000种,约占总维管束植物种类的10%。林冠附生物的生物量在世界各地森林中存在较大的差异,其范围在105~44 000 kg·hm^-2之间,其中在一些热带和温带天然老龄林中林冠附生物的生物量超过了宿主林木的叶生物量。林冠附生植物还具有较大的叶面积指数(LAI)。林冠附生物丰富的物种组成、较高的生物量、独特的生理形态特征以及它们分布于森林与大气相互作用的关键界面,使得它们在生态系统物种多样性形成及其维持机制、养分和水分循环、指示环境质量等方面具有重要的作用。林冠附生植物及其枯死残留物具有较强的能力吸收雨水和空气中的营养物质,在林冠层中形成一个潮湿的环境促进氮固定,林冠附生植物群落还能为生存于林冠的其它生物(如鸟类、哺乳动物、两栖动物、爬行动物和昆虫等)提供食物和栖息场所。林冠附生植物的生长发育与分布格局除与宿主有关外,还受到环境因素(气候、地形、微生境条件等)和人为干扰的影响。由于世界各地森林类型多样和环境条件各异,目前国际上有关附生物的研究仍十分活跃,建立了林冠研究网络,研究不同类型森林中附生植物及其枯死残留物的动态及其与群落特征、环境因子的数量关系,探讨、交流和发展有效的标准测量方法和技术是该领域研究的主要内容。国内对林冠附生植物生态学的研究刚刚起步,有待于今后加强该领域的研究。     

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.     

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.