Small plants, large plants: the importance of plant size for the physiological ecology of vascular epiphytes

Recently, a number of publications have reported that many physiological properties of vascular epiphytes are a function of plant size. This short review will summarize what is known to date about this phenomenon, describe the possible mechanism and will discuss the consequences for the present understanding of epiphyte biology. Size-related changes are also known from other plant groups and it is argued that close attention should be paid to the size of the organisms under study in order to understand the performance and survival of a species in the field. In the light of these findings, the results of many earlier studies on epiphyte ecophysiology are now difficult to interpret because essential information on the size of the specimens used is missing.     

Vascular epiphytes in the temperate zones–a review

Vascular epiphytes are typically associated with tropical rainforests, whereas their occurrence in temperate forests is little appreciated. This review summarises the available information on epiphytism in the temperate zones (> 23.5 latitude), which has not been reviewed omprehensively for more than a century, and critically analyses the proposed mechanisms behind the observed biogeographical patterns. Although in the temperate zone epiphytic vascular plants are rarely as impressive as in tropical forests, there are noteworthy exceptions. Temperate rain forests of Chile and New Zealand, or montane forests in the Himalayas are comparable to many tropical forests in terms of epiphyte biomass and diversity, but differ in their taxonomic spectrum temperate epiphyte communities are generally dominated by ferns and fern-allies. Other temperate areas are not, however, necessarily barren of epiphytes, as repeatedly implied. Quite in contrast, local populations of epiphytes in a large number of other non-tropical areas in both the southern and the northern hemisphere can be quite conspicuous. The proposed reasons for the latitudinal gradients in epiphyte abundance and diversity (water scarcity or low tempera-tures). are not fully convincing and, moreover, still await experimental verification. Other factors, both historical (e.g., Pleistocene extinctions) and ecological (e.g., prevalence of conifers in the northern hemisphere), should also be taken into consideration to obtain a comprehensive explanation of the extant global distribution of vascular epiphytes.     

The systematic distribution of vascular epiphytes – a critical update

Vascular epiphyte species exclusively, or at least primarily, germinate and grow on other plants without contact with the soil and, in contrast to mistletoes, without parasitizing their hosts. The last review of the systematic distribution of this diverse group of plants dates back more than two decades. The present study pursues three major goals: (1) it critically discusses conceptual problems arising from the distinction of epiphytes from non‐epiphytes; (2) it presents a compilation of epiphytic diversity derived from a vast number of sources; and (3) it arranges epiphyte diversity in an up‐to‐date taxonomic framework. The resulting compilation, which identifies 27 614 species of vascular epiphytes (including primary hemiepiphytes) representing 913 genera in 73 families, or approximately 9% of extant vascular plant diversity, is meant to be an important tool for studies on the ecology and evolution of epiphytes, but also for comparative studies with a focus on other life forms. © 2013 The Linnean Society of London     

中国森林附生维管植物多样性数据集

附生维管植物是热带、亚热带湿性山地森林植物群落中物种最为丰富的类群之一, 在维系森林生物多样性及生态系统功能与服务中发挥着重要作用, 然而附生维管植物调查相对困难, 在多样性调查和统计中往往被忽视, 迄今为止我国尚缺乏全国范围的森林附生维管植物名录。本项研究基于已公开发布的数据库, 搜集并整理了1982-2022年间的关于中国森林附生维管植物研究的文献书籍。首先, 提取文献资料中的有效信息, 依据物种2000中国节点的数据进行标准化校正, 整理出中国森林附生维管植物共计49科246属1,739种, 然后据此编写了相对准确、完整的中国森林附生维管植物物种名录。本名录作为我国第一个全国性森林附生维管植物名录, 可为森林生物多样性保护工作提供重要的数据支持, 并能为深入研究附生维管植物的进化生物学、生物地理学及宏观生态学等领域提供重要的基础信息。     

Agent‐based modeling of the effects of forest dynamics,selective logging,and fragment size on epiphyte communities

1. Forest canopies play a crucial role in structuring communities of vascular epiphytes by providing substrate for colonization, by locally varying microclimate, and by causing epiphyte mortality due to branch or tree fall. However, as field studies in the three‐dimensional habitat of epiphytes are generally challenging, our understanding of how forest structure and dynamics influence the structure and dynamics of epiphyte communities is scarce.
2. Mechanistic models can improve our understanding of epiphyte community dynamics. We present such a model that couples dispersal, growth, and mortality of individual epiphytes with substrate dynamics, obtained from a three‐dimensional functional–structural forest model, allowing the study of forest–epiphyte interactions. After validating the epiphyte model with independent field data, we performed several theoretical simulation experiments to assess how (a) differences in natural forest dynamics, (b) selective logging, and (c) forest fragmentation could influence the long‐term dynamics of epiphyte communities.
3. The proportion of arboreal substrate occupied by epiphytes (i.e., saturation level) was tightly linked with forest dynamics and increased with decreasing forest turnover rates. While species richness was, in general, negatively correlated with forest turnover rates, low species numbers in forests with very‐low‐turnover rates were due to competitive exclusion when epiphyte communities became saturated. Logging had a negative impact on epiphyte communities, potentially leading to a near‐complete extirpation of epiphytes when the simulated target diameters fell below a threshold. Fragment size had no effect on epiphyte abundance and saturation level but correlated positively with species numbers.
4. Synthesis: The presented model is a first step toward studying the dynamic forest–epiphyte interactions in an agent‐based modeling framework. Our study suggests forest dynamics as key factor in controlling epiphyte communities. Thus, both natural and human‐induced changes in forest dynamics, for example, increased mortality rates or the loss of large trees, pose challenges for epiphyte conservation.

     

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.     

Seedling establishment of vascular epiphytes on isolated and enclosed forest trees in an Andean landscape, Ecuador

The impact of human disturbance on colonisation dynamics of vascular epiphytes is poorly known. We studied abundance, diversity and floristic composition of epiphyte seedling establishing on isolated and adjacent forest trees in a tropical montane landscape. All vascular epiphytes were removed from plots on the trunk bases of Piptocoma discolor. Newly established epiphyte seedlings were recorded after 2 years, and their survival after another year. Seedling density, total richness at family and genus level, and the number of families and genera per plot were significantly reduced on isolated trees relative to forest trees. Seedling assemblages on trunks of forest trees were dominated by hygrophytic understorey ferns, those on isolated trees by xerotolerant canopy taxa. Colonisation probability on isolated trees was significantly higher for plots closer to forest but not for plots with greater canopy or bryophyte cover. Seedling mortality on isolated trees was significantly higher for mesophytic than for xerotolerant taxa. Our results show that altered recruitment can explain the long-term impoverishment of post-juvenile epiphyte assemblages on isolated remnant trees. We attribute these changes to a combination of dispersal constraints and the harsher microclimate documented by measurements of temperature and humidity. Although isolated trees in anthropogenic landscapes are considered key structures for the maintenance of forest biodiversity in many aspects, our results show that their value for the conservation of epiphytes can be limited. We suggest that abiotic seedling requirements will increasingly constitute a bottleneck for the persistence of vascular epiphytes in the face of ongoing habitat alteration and atmospheric warming.     

“No signs of saturation”: long-term dynamics of vascular epiphyte communities in a human-modified landscape

It is debated whether slowing human population growth and intense urbanization may result in a slowdown of deforestation and an acceleration of natural forest regeneration. In a fragmented landscape the structure and composition of developing secondary forests will strongly depend on the local species pool. Thus, the understanding how organisms cope with biotic and abiotic challenges outside pristine habitats is pivotal. Structurally dependent, vascular epiphytes are an important biotic component of tropical forests. In human-modified tropical landscapes potential hosts are often still present. We aimed to assess if human-modified landscapes offer a refuge habitat in which epiphytes can form metacommunities that are ultimately viable. Eight years after an initial assessment we recensused the epiphytes in pasture trees in western Panama along a strong rainfall gradient. We document a threefold abundance increase (ca. 20,000 vs. >60,000 individuals) and an increase in species-richness (66 vs. 86 mostly drought-tolerant species). This large net increase suggests a highly dynamic system. Although absolute abundances changed dramatically, the relative contribution of major taxonomic groups to overall diversity and abundance changed little. Neither rainfall nor tree growth had a significant effect on relative annual community growth rates. At the plot level (=metacommunities), abundance increase was mostly due to the species already present in the first census, at the tree level (=communities) the contribution of new and old species was comparable. The documented long-term trend in epiphyte metacommunities in a human-modified landscape suggests that a diverse set of species sustains viable metacommunities and is likely to provide structural diversity to developing secondary forest.     

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.     

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 Conservation Value of Secondary Forests for Vascular Epiphytes in Central Panama

Secondary forests that develop following land abandonment could compensate for the losses of diversity and structure that accompany deforestation of old‐growth forests in tropical regions. Whether secondary forests can harbor similar species richness, density, and composition of old‐growth forests for vascular epiphytes remains largely unknown for secondary forests older than 50 yr. We examined community structure (species richness, density, and species composition) of vascular epiphytes in older secondary forests between 35 and 115 yr after land abandonment and nearby old‐growth forests to determine if the community structure of epiphytes in secondary forests approaches that of old‐growth forests over time. The recovery of epiphyte species richness was rapid with 55‐year‐old forests containing 65 percent of old‐growth epiphyte species richness. Secondary forest epiphyte communities were found to be statistically nested within secondary forests older in age and within old‐growth forests. Similarity of epiphyte communities to old‐growth forests increased to 75 percent, 115 yr after abandonment. This study suggests that secondary forests will likely recover old‐growth epiphyte richness and composition given enough time. Epiphyte densities did not recover quickly with 55‐year‐old forests having 14 percent and 115‐year‐old forests having only 49 percent of the density of old‐growth forest epiphytes. The low density of epiphytes in secondary forests could impact rainforest diversity and function. We conclude that in less than 115 yr, although secondary moist forests have high conservation value for some aspects of community structure, they are unlikely to compensate biologically for the loss of diversity and ecosystem function that high epiphyte densities provide.     

Epiphytic (including hemiepiphytes) diversity in three timber species in the southwestern Amazon, Brazil

Forestry managers have been searching for ways to reduce the impacts of logging on Amazonian biodiversity, but some basic factors are still not considered in native forestry operations, among them the diversity of epiphytes associated with the logged trees. Our goals in this study were to determine the floristic composition, quantify the species richness, and characterize the species diversity of the vascular epiphytic community present in three timber tree species in Acre State, Brazil. We collected and identified all epiphytes in 30 randomly selected trees ≥35 cm DBH of each of three important timber species, Tabebuia serratifolia, Manilkara inundata and Couratari macrosperma. We also documented the epiphyte diversity in 120 randomly selected trees ≥35 cm DBH of 56 other species to determine whether the three timber species have different epiphyte diversity than the tree community at large. The epiphyte samples in the three timber species showed 77 species, 13 of which were new records for the flora of Acre state. The epiphyte community in the randomly selected trees presented a total of 56 species. The timber species phorophytes hosted on average three times more epiphyte species per tree than the other 120 randomly selected trees. These results show that a substantial portion of local floristic richness can be lost during logging activity due if not properly managed by rescuing epiphytes after felling the trees. Although these epiphytes could contribute positively to forestry sustainability due to their ornamental value, increasing the economic yield per hectare, there are no local initiatives for economic use of epiphytes.     

DISTRIBUTION OF ALGAL EPIPHYTES ACROSS ENVIRONMENTAL GRADIENTS AT DIFFERENT SCALES: INTERTIDAL ELEVATION,HOST CANOPIES,AND HOST FRONDS1

Understanding epiphyte distribution in coastal communities is important because these organisms affect many others directly or indirectly. Yet, their distribution has been considerably less studied than that of their hosts and other primary‐space holders. Identifying major sources of variation in epiphyte abundance is thus still a need. Environmental gradients help predict species distribution and are pervasive on marine shores. In this study, we test the notion that environmental gradients across intertidal elevation, throughout host canopies, and along host fronds explain a large variation in the abundance of sympatric epiphytes. Our model system was the assemblage of Ascophyllum nodosum (L.) Le Jol. and its epiphytes Vertebrata lanosa (L.) T. A. Chr. [= Polysiphonia lanosa (L.) Tandy], Elachista fucicola (Velley) Aresch., and Pylaiella littoralis (L.) Kjellm. On the coast of Nova Scotia (Canada), we found evidence of a spatial segregation among these species at almost all scales. While the red epiphyte V. lanosa was more common at high‐ and midintertidal elevations (peaking at midelevations) and on middle segments of host fronds, the brown epiphytes E. fucicola and P. littoralis were more common at low elevations and restricted to distal segments of host fronds. Canopy habitat affected abundance only for V. lanosa, which was more common within the host canopy than on its periphery at midelevations. Since the studied gradients are related to predictable changes in abiotic factors, the identification of likely causes behind the observed patterns was facilitated. Our study ends by proposing abiotic and biotic factors that deserve priority in the experimental testing of the forces structuring this assemblage.     

Vascular epiphyte populations with higher leaf nutrient concentrations showed weaker resilience to an extreme drought in a montane cloud forest

• Leaf stoichiometry can characterize plant ecological strategies and correlate with plant responses to climate change. The role of vascular epiphytes in the ecosystem processes of tropical and subtropical forest ecosystems cannot be ignored. Vascular epiphytes are very vulnerable to climate change, however, the relationship between the response of epiphytes to climate change and leaf stoichiometry is not well understood.
• We present data for 19 vascular epiphyte species that were collected during four consecutive censuses (in 2005, 2010, 2015, and 2020) over 15 years in a subtropical montane cloud forest. We assessed the relationships between the population dynamics and leaf stoichiometry of these vascular epiphytes.
• Experiencing an extreme drought, 14 of the 19 epiphyte species showed an obvious decrease in the number of individuals, and all species showed negative growth in the number of populations. Subsequently, the total number of individuals gradually recovered, increasing from 7,195 in 2010 to 10,121 in 2015, then to 13,667 in 2020. The increase in the number of vascular epiphyte individuals from 2010 to 2015 was significantly negatively correlated with leaf nitrogen and phosphorus concentration, and was significantly positively correlated with the leaf carbon-nitrogen ratio.
• Vascular epiphyte populations with higher leaf nutrient concentrations exhibited weaker resilience to the extreme drought, which demonstrated that a resource-conservative strategy was advantageous for the recovery of epiphyte populations. Our findings suggest that ecological stoichiometry can be a useful framework for forecasting the dynamics of vascular epiphyte populations in response to climate change.

     

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.     

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.     

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。     

Interaction network of vascular epiphytes and trees in a subtropical forest

The commensalistic interaction between vascular epiphytes and host trees is a type of biotic interaction that has been recently analysed with a network approach. This approach is useful to describe the network structure with metrics such as nestedness, specialization and interaction evenness, which can be compared with other vascular epiphyte-host tree networks from different forests of the world. However, in several cases these comparisons showed different and inconsistent patterns between these networks, and their possible ecological and evolutionary determinants have been scarcely studied. In this study, the interactions between vascular epiphytes and host trees of a subtropical forest of sierra de San Javier (Tucuman, Argentina) were analysed with a network approach. We calculated metrics to characterize the network and we analysed factors such as the abundance of species, tree size, tree bark texture, and tree wood density in order to predict interaction frequencies and network structure. The interaction network analysed exhibited a nested structure, an even distribution of interactions, and low specialization, properties shared with other obligated vascular epiphyte-host tree networks with a different assemblage structure. Interaction frequencies were predicted by the abundance of species, tree size and tree bark texture. Species abundance and tree size also predicted nestedness. Abundance indicated that abundant species interact more frequently; and tree size was an important predictor, since larger-diameter trees hosted more vascular epiphyte species than small-diameter trees. This is one of the first studies analyzing interactions between vascular epiphytes and host trees using a network approach in a subtropical forest, and taking the whole vascular epiphyte assemblage of the sampled community into account.     

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.     

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.