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Background and ScopeThe epiphytic life form characterizes almost 10 % of all vascular plants. Defined by structural dependence throughout their life and their non-parasitic relationship with the host, the term epiphyte describes a heterogeneous and taxonomically diverse group of plants. This article reviews the importance of crassulacean acid metabolism (CAM) among epiphytes in current climatic conditions and explores the prospects under global change.Results and ConclusionsWe question the view of a disproportionate importance of CAM among epiphytes and its role as a ‘key innovation’ for epiphytism but do identify ecological conditions in which epiphytic existence seems to be contingent on the presence of this photosynthetic pathway. Possibly divergent responses of CAM and C3 epiphytes to future changes in climate and land use are discussed with the help of experimental evidence, current distributional patterns and the results of several long-term descriptive community studies. The results and their interpretation aim to stimulate a fruitful discussion on the role of CAM in epiphytes in current climatic conditions and in altered climatic conditions in the future.

Vascular epiphytes represent almost 10% of all terrestrial plant diversity. Despite the extensive research on the functional ecology and challenges of epiphytic growth, there is still very little known on how exposure to mechanically induced stress affects the growth and development of epiphytes. Therefore, this study investigated the effect of such mechanical stress on the growth and biomass allocation of epiphytic bromeliads. Juvenile plants of two species were subjected to two types of mechanical stress in the greenhouse—permanent displacement and temporary, recurring mechanical flexing. ANOVAs were used to test possible treatment effects on growth, root–shoot ratio, root diameter, and root area distribution ratio. Contrary to previous studies on herbaceous plants, these bromeliads showed little to no change in root and shoot properties in either species. The root–shoot ratio increased in disturbedGuzmania lingulataplants, but not inVrieseasp. Treatment effects on growth were inconsistent: a stress effect on growth was significant only in the first 2 months of the experiment inG. lingulata, whilst none of the stress treatments negatively affected growth inVrieseasp. All disturbed plants showed some degree of curvature on their stems and leaves against the area of stress to obtain an upright position. This was probably related to the maintenance of a functional tank. This study provides quantitative and qualitative data to understand thigmomorphogenic responses of bromeliads to mechanical stress. Future studies could include field surveys to quantify on-site mechanical stresses and the corresponding morphological changes in vascular epiphytes.

For vascular epiphytes, secure attachment to their hosts is vital for survival. Yet studies detailing the adhesion mechanism of epiphytes to their substrate are scarce. Examination of the root hair-substrate interface is essential to understand the attachment mechanism of epiphytes to their substrate. This study also investigated how substrate microroughness relates to the root-substrate attachment strength and the underlying mechanism(s).Seeds of Anthurium obtusum were germinated, and seedlings were transferred onto substrates made of epoxy resin with different defined roughness. After 2 months of growth, roots that adhered to the resin tiles were subjected to anchorage tests, and root hair morphology at different roughness levels was analyzed using light and cryo scanning electron microscopy.The highest maximum peeling force was recorded on the smooth surface (glass replica, 0 µm). Maximum peeling force was significantly higher on fine roughness (0, 0.3, 12 µm) than on coarse (162 µm). Root hair morphology varied according to the roughness of the substrate. On smoother surfaces, root hairs were flattened to achieve large surface contact with the substrate. Attachment was mainly by adhesion with the presence of a glue-like substance. On coarser surfaces, root hairs were tubular and conformed to spaces between the asperities on the surface. Attachment was mainly via mechanical interlocking of root hairs and substrate.This study demonstrates for the first time that the attachment mechanism of epiphytes varies depending on substrate microtopography, which is important for understanding epiphyte attachment on natural substrates varying in roughness.

1. Plant functional traits impact the fitness and environmental niche of plants. Major plant functional types have been characterized by their trait spectrum, and the environmental and phylogenetic imprints on traits have advanced several ecological fields. Yet, very few trait data on epiphytes, which represent almost 10% of vascular plants, are available. 2. We collated 76,561 trait observations for 2,882 species of vascular epiphytes and compared these to non-epiphytic herbs and trees to test hypotheses related to how the epiphytic habit affects traits, and if epiphytes occupy a distinct region in the global trait space. We also compared variation in traits among major groups of epiphytes, and investigated the coordination of traits in epiphytes, ground-rooted herbs and trees. 3. Epiphytes differ from ground-rooted plants mainly in traits related to water relations. Unexpectedly, we did not find lower leaf nutrient concentrations, except for nitrogen. Mean photosynthetic rates are much lower than in ground-rooted plants and lower than expected from the nitrogen concentrations. Trait syndromes clearly distinguish epiphytes from trees and from most non-epiphytic herbs. 4. Among the three largest epiphytic taxa, orchids differ from bromeliads and ferns mainly by having smaller and more numerous stomata, while ferns differ from bromeliads by having thinner leaves, higher nutrient concentrations, and lower water content and water use efficiency. 5. Trait networks differ among epiphytes, herbs and trees. While all have central nodes represented by SLA and mass-based photosynthesis, in epiphytes, traits related to plant water relations have stronger connections, and nutrients other than potassium have weaker connections to the remainder of the trait network. Whereas stem-specific density reflects mechanical support related to plant size in herbs and trees, in epiphytes it mostly reflects water storage and scales with leaf water content. 6. Synthesis. Our findings advance our understanding of epiphyte ecology, but we note that currently mainly leaf traits are available. Important gaps are root, shoot and whole plant, demographic and gas exchange traits. We suggest how future research might use available data and fill data gaps. ISSN:0022-0477

Vascular epiphytes represent almost 10% of all terrestrial plant diversity. Despite their growing sites exposed to frequent extensive storms, nothing is known about how external mechanical perturbations by wind affects epiphytes’ growth and development. Therefore, this study investigated the effect of such mechanical stress on the growth rate and morphology of epiphytes. Juvenile plants of two species of bromeliads, grown in the greenhouse, were subjected to varying degrees of mechanical stress – Permanent displacement, tilting and mechanical flexing. ANOVAs were used to test possible treatment effects on growth, root-shoot ratio, root diameter, and leeward-windward root area ratio. Contrary to previous studies on herbaceous plants, these bromeliads showed little to no change in root and shoot properties in either species. Although the root-shoot ratio increased in disturbed Guzmania lingulata plants (Tukey HSD, p < 0.001), it was not observed in Vriesea sp. Treatment effects on growth were inconsistent: stress effect on growth was significant only in the first three months of the experiment (padj < 0.05) in G. lingulata, while none of the stress treatments resulted in negative growth in Vriesea sp. All disturbed plants showed some degree of curvature on their stems and leaves against the area of stress to obtain an upright position. This was probably related to the maintenance of a functional tank. This study provided quantitative and qualitative data to understand thigmomorphogenic responses of bromeliads to mechanical stress. Future studies could include field surveys to quantify on-site wind stress and the corresponding morphological changes in vascular epiphytes.

Vascular epiphytes represent almost 10% of all terrestrial plant diversity. Being structurally dependent on trees, epiphytes live at the interface of vegetation and atmosphere, making them susceptible to atmospheric changes. Despite the extensive research on vascular epiphytes, little is known about wind disturbance on these plants. Therefore, this study investigated the wind-epiphyte mechanical interactions by quantifying the drag forces on epiphytic bromeliads when subjected to increasing wind speeds (5-22 m s-1) in a wind tunnel. Drag coefficients (Cd) and Vogel exponents (B) were calculated to quantify the streamlining ability of different bromeliad species. Bromeliads' reconfiguration occurred first via bending and aligning leaves in the flow direction. Then leaves clustered and reduced the overall plant frontal area. This reconfiguration caused drag forces to increase at a slower rate as wind velocity increased. In the extreme case, drag force was reduced by 50% in a large Guzmania monostachia individual at a wind velocity of 22 m s-1, compared to a stiff model. This species had one of the smallest Cd (0.58) at the highest wind velocity, and the largest negative mean B (-0.98), representing the largest reconfiguration capacity amongst the tested bromeliads. The streamlining ability of bromeliads was mainly restricted by the rigidity of the lower part of the plant where the leaves are already densely clustered. Wind speeds used in this study were generally low as compared to storm force winds. At these low wind speeds, reconfiguration was an effective mechanism for drag reduction in bromeliads. This mechanism is likely to lose its effectiveness at higher wind speeds when continuous vigorous fluttering results in leaf damage and aspects such as root-attachment strength and substrate stability become more relevant. This study is a first step towards an understanding of the mechanical bottleneck in the epiphyte-tree-system under wind stress.

Vascular epiphytes represent almost 10% of all terrestrial plant diversity. Being structurally dependent on trees, epiphytes live at the interface of vegetation and atmosphere, making them susceptible to atmospheric changes. Despite the extensive research on vascular epiphytes, little is known about wind disturbance on these plants. Therefore, this study investigated the wind-epiphyte mechanical interactions by quantifying the drag forces on epiphytic bromeliads when subjected to increasing wind speeds (5–22 m s-1) in a wind tunnel. Drag coefficients (Cd) and Vogel exponents (B) were calculated to quantify the streamlining ability of different bromeliad species. Bromeliads’ reconfiguration occurred first via bending and aligning leaves in the flow direction. Then leaves clustered and reduced the overall plant frontal area. This reconfiguration caused drag forces to increase at a slower rate as wind velocity increased. In the extreme case, drag force was reduced by 50% in a large Guzmania monostachia individual at a wind velocity of 22 m s-1, compared to a stiff model. This species had one of the smallest Cd (0.58) at the highest wind velocity, and the largest negative mean B (-0.98), representing the largest reconfiguration capacity amongst the tested bromeliads. The streamlining ability of bromeliads was mainly restricted by the rigidity of the lower part of the plant where the leaves are already densely clustered. Wind speeds used in this study were generally low as compared to storm force winds. At these low wind speeds, reconfiguration was an effective mechanism for drag reduction in bromeliads. This mechanism is likely to lose its effectiveness at higher wind speeds when continuous vigorous fluttering results in leaf damage and aspects such as root-attachment strength and substrate stability become more relevant. This study is a first step towards an understanding of the mechanical bottleneck in the epiphyte-tree-system under wind stress.

Root morphology is expected to respond to environmental conditions. Two earlier studies reported predictable changes in the structure of the velamen radicum (common in monocotyledons) along rainfall gradients, which was expected from its assumed role in plant-water-relations. The present study expanded on this idea by analysing nine root traits that can be linked to root function, including velamen structure, along two moisture gradients of different spatial scales: 1) along the vertical gradient of a lowland forest in central Panama and 2) along a strong regional rainfall gradient in western Panama. All studied traits (e.g. velamen width, number of xylem poles, lignification of the exodermis) of the 45 orchid species showed substantial intraspecific and interspecific variation, but none of this (within forest) or very little of this (regional gradient) was related to the gradients. Only the community weighted means of velamen width, stele width and xylem pole number varied significantly with rainfall, with substantial effect sizes only found in the latter two traits. Our results clearly disagree with those of earlier publications, which did not include species abundance and/or rigorous statistics in their analyses. This discrepancy highlights that our understanding of the link between velamen form and function is still rudimentary.

Although it has long been acknowledged that seed characteristics are of major importance to the conquest of tree crowns by vascular epiphytes, there is surprisingly little quantitative evidence on the aerodynamic properties of their diaspores. We used a custom-built device to determine the terminal velocity (Vterm) of falling seeds, a parameter that has been shown to have high predictive power for the wind dispersal potential of diaspores under natural conditions. We determined Vterm of 45 species of epiphytic and terrestrial Orchidaceae, which almost doubles the currently available database for this family. Although varying by a factor of five with values of 0.09–0.4 m s−1, Vterm was invariably very slow compared to plants in general. For each species, we also took morphological data and determined seed mass. None of these parameters was linearly correlated with Vterm and neither did the average Vterm differ between species found in the two habitats, although seeds of terrestrial taxa were significantly larger and heavier. Finally, we demonstrate the potential of our device to measure Vterm of even smaller diaspores by successfully quantifying Vterm of fern spores. This tool has much potential for the quantitative study of dispersal of plants with tiny diaspores, particularly in a conservation context in fragmented landscapes.

Tank bromeliads are prominent and distinctive components of many Neotropical forests. Functionally important for the plants themselves these phytotelms also represent important freshwater habitats in the forest canopy for a range of animal species and other biota. Existing literature values of the amount of water held in these tanks have never been systematically compiled nor has there ever been a systematic assessment of the water holding capacity of bromeliad tanks (tank capacity, TC) with a consistent methodology that covered a large taxonomic breadth of the bromeliad family. We determined TC for more than 200 taxa and collected additional data on total plant dry mass, projected area of the rosette (catchment area), and the number of individual reservoirs for a subset of them. These data were then used to establish intraspecific and interspecific allometric relationships of TC with plant size, the water held in living tissue, and the amount of rain necessary to fill the tank. This paper provides important baseline data for any study of tank function and gives a more modest picture of the “typical” bromeliad tank as often implied by the reference to extreme and unsubstantiated values in the literature.

Background: An ever-increasing proportion of tropical ecosystems are affected by on-going land-use changes, stressing the importance of understanding how organisms cope with biotic and abiotic challenges outside their natural habitat. An important group in the tropics are vascular epiphytes, whose response to human disturbance is poorly understood. Aims: Creating a baseline data set of epiphyte diversity in human settlements and assessing differences with assemblages of less disturbed habitats. Methods: We surveyed the vascular epiphyte assemblages in 25 settlements of south-west Panama along a rainfall gradient. We tested how epiphyte assemblages were affected by human disturbance and climate by comparing our data to that of pastures and forest. Results: Almost half (238 of 499) of all studied trees hosted at least one epiphyte. Altogether, 10,700 epiphytes of 56 species were found. Rainfall strongly affected both composition and diversity of epiphyte assemblages, whereas geographic proximity did not. In settlements, the species pool was considerably smaller than in pastures and undisturbed forest; however, settlements’ diversity was only significantly lower compared to pastures. Conclusions: Epiphyte assemblages in settlements had a diminished species pool but diversity per tree was comparable to less disturbed habitats. Unsurprisingly, rainfall seems to be the main determinant of epiphyte diversity in rural settlements.

The hosts of vascular epiphytes differ in many ways, not least in leaf phenology. We hypothesized that differences in microclimatic conditions in evergreen vs. deciduous trees would affect epiphytes at various levels, from organ physiology to community structure. Indeed, deciduous tree species hosted less abundant and species-poorer epiphyte assemblages. Physiologically, epiphyte assemblages differed in the proportion of CAM species and individuals, and in SLA and δ13C values. Effects were also detectable at a demographic level, i.e. in growth and survival rates. Although not all studied epiphyte species showed these effects, the data generally support our basic hypothesis., The processes that govern diverse tropical plant communities have rarely been studied in life forms other than trees. Structurally dependent vascular epiphytes, a major part of tropical biodiversity, grow in a three-dimensional matrix defined by their hosts, but trees differ in their architecture, bark structure/chemistry and leaf phenology. We hypothesized that the resulting seasonal differences in microclimatic conditions in evergreen vs. deciduous trees would affect epiphytes at different levels, from organ physiology to community structure. We studied the influence of tree leaf phenology on vascular epiphytes on the Island of Barro Colorado, Panama. Five tree species were selected, which were deciduous, semi-deciduous or evergreen. The crowns of drought-deciduous trees, characterized by sunnier and drier microclimates, hosted fewer individuals and less diverse epiphyte assemblages. Differences were also observed at a functional level, e.g. epiphyte assemblages in deciduous trees had larger proportions of Crassulacean acid metabolism species and individuals. At the population level a drier microclimate was associated with lower individual growth and survival in a xerophytic fern. Some species also showed, as expected, lower specific leaf area and higher δ13C values when growing in deciduous trees compared with evergreen trees. As hypothesized, host tree leaf phenology influences vascular epiphytes at different levels. Our results suggest a cascading effect of tree composition and associated differences in tree phenology on the diversity and functioning of epiphyte communities in tropical lowland forests.