A new approach to an old problem: how to categorize the habit of ferns and lycophytes.

BACKGROUND AND AIMS: Substrate preferences are often treated as species traits and are used to distinguish different habits, i.e. an epiphytic, lithophytic or terrestrial habit. Such a categorization, however, ignores substantial intraspecific variation. An approach that takes biological variability within a species into account is needed. METHODS: We focused on four large genera of ferns and lycophytes and found relevant information in >500 sources, such as online databases, checklists, floras and species descriptions. Translating textual information into a quantitative index, we quantified the propensity to grow on either substrate as a continuous trait for 1475 species. KEY RESULTS: Only a minority of species exhibited strict substrate fidelity, but a majority of them showed clear habitat preferences. The relative frequencies of intermediates between strict lithophytes, epiphytes and terrestrials does not support the frequent notion of ecological similarity of the lithophytic and epiphytic habitat. CONCLUSIONS: The compiled data are useful immediately for ecological and evolutionary studies with the focal taxa. More importantly, we propose the replacement of the concept of distinct habits with one of gradual differences. This should have a profound impact on any such study with plants in general.

Does A Velamen Radicum Effectively Protect Epiphyte Roots against Excessive Infrared Radiation?

Velamen radicum, a dead tissue at maturity, characterizes the roots of many epiphytes. Apart from a role in water and nutrient uptake, protection against excessive radiation in the upper forest canopy has also been suggested, but this function has never been critically assessed. To test this notion, we studied the roots of 18 species of Orchidaceae and Araceae. We defined thermal insulation traits of velamina by monitoring the temperature on the velamen surface and just below the velamen while exposing it to infrared radiation. We investigated velamen's functionality-correlating morphological and thermal insulation traits. In addition, we investigated the viability of the living root tissue after heat exposure. The maximal surface temperatures ranged from 37-51 °C, while the temperature difference between the upper and lower velamen surface (i.e., ∆Tmax) ranged from 0.6-3.2 °C. We found a relationship of velamen thickness with ∆Tmax. Tissue viability was strongly affected by temperatures >42 °C, and no significant recovery after heat exposure was found. Thus, there is only limited support for an insulating function of velamen, but the data suggest considerable species-specific differences in heat tolerance. The latter could be a crucial determinant of the vertical distribution of epiphytes.

Smoothing out the misconceptions of the role of bark roughness in vascular epiphyte attachment.

Vascular epiphytes represent c. 10% of all vascular plant species. In epiphytes, attachment is essential for survival throughout consecutive ontogenetic stages of their life, starting with: (1) initial propagule attachment to the host; followed by (2) the development of first root-substrate connections; and (3) maintenance of this attachment despite increased size and mechanical disturbances by rain, wind, or crossing animals. Although structural dependence on a host is a defining characteristic of an epiphyte, the fundamental mechanism(s) of how these plants initially attach and remain attached to their hosts remain poorly understood. Bark characteristics such as stability and roughness have been highlighted as keys to an understanding of this connection. Here, we stress that the understanding of how an epiphyte attaches itself to the substrate is central for a meaningful quantification and interpretation of bark roughness. Without explicit information on the attachment mechanism or the relative sizes of the attaching structures, simply linking a haphazardly chosen index of bark roughness to epiphyte establishment is flawed. This review introduces a conceptual framework to explain the mechanistic link between epiphytes and host in different ontogenetic stages and should guide future work designed to improve our understanding of this vital part of epiphyte ecology.

CAM plants: their importance in epiphyte communities and prospects with global change.

BACKGROUND AND SCOPE: The 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 CONCLUSIONS: We 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.

New Proposal of Epiphytic Bromeliaceae Functional Groups to Include Nebulophytes and Shallow Tanks.

The Bromeliaceae family has been used as a model to study adaptive radiation due to its terrestrial, epilithic, and epiphytic habits with wide morpho-physiological variation. Functional groups described by Pittendrigh in 1948 have been an integral part of ecophysiological studies. In the current study, we revisited the functional groups of epiphytic bromeliads using a 204 species trait database sampled throughout the Americas. Our objective was to define epiphytic functional groups within bromeliads based on unsupervised classification, including species from the dry to the wet end of the Neotropics. We performed a hierarchical cluster analysis with 16 functional traits and a discriminant analysis, to test for the separation between these groups. Herbarium records were used to map species distributions and to analyze the climate and ecosystems inhabited. The clustering supported five groups, C3 tank and CAM tank bromeliads with deep tanks, while the atmospheric group (according to Pittendrigh) was divided into nebulophytes, bromeliads with shallow tanks, and bromeliads with pseudobulbs. The two former groups showed distinct traits related to resource (water) acquisition, such as fog (nebulophytes) and dew (shallow tanks). We discuss how the functional traits relate to the ecosystems inhabited and the relevance of acknowledging the new functional groups.

Holding on or falling off: The attachment mechanism of epiphytic Anthurium obtusum changes with substrate roughness.

PREMISE: 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). METHODS: 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. RESULTS: 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. CONCLUSIONS: 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.

Go with the flow: The extent of drag reduction as epiphytic bromeliads reorient in wind.

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.

Dispersal and establishment of vascular epiphytes in human-modified landscapes.

The ongoing destruction of old-growth forests puts tropical forest species under great pressure because of the resulting habitat loss. A pre-requisite for the maintenance of a viable metacommunity in a fragmented landscape is the connectivity between habitable patches. We experimentally studied four vital steps of epiphyte dispersal in different habitat types in western Panama. (i) Seed falling velocity (Vterm) is known to correlate with long-distance dispersal via convective updraft. All measured Vterm of bromeliad and orchid seeds fell into a range of velocities with a high chance of long-distance dispersal. (ii) We quantified attachment success of bromeliad seeds as a function of bark rugosity with >30 common tree species in the region. Even fine bark structure allowed effective attachment. (iii and iv) Successful establishment is achieved when a seed germinates and a plantlet grows and survives. Germination success and early establishment of four bromeliad species did not differ between isolated trees, teak plantations or secondary forest patches. Microclimatic differences between habitat types were marginal and neither germination nor establishment correlated significantly with annual precipitation. The findings suggest a large capacity for dispersal and successful early establishment for these anemochorous species. A potentially regenerating forest may receive considerable input from sources such as pasture trees and in this way gain structural complexity, which also greatly enhances its value for other forest organisms.

Host tree phenology affects vascular epiphytes at the physiological, demographic and community level.

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 δ(13)C 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.