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Epiphytes make up roughly 10% of all vascular plant species globally and play important functional roles, especially in tropical forests. However, to date, there is no comprehensive list of vascular epiphyte species. Here, we present EpiList 1.0, the first global list of vascular epiphytes based on standardized definitions and taxonomy. We include obligate epiphytes, facultative epiphytes, and hemiepiphytes, as the latter share the vulnerable epiphytic stage as juveniles. Based on 978 references, the checklist includes >31,000 species of 79 plant families. Species names were standardized against World Flora Online for seed plants and against the World Ferns database for lycophytes and ferns. In cases of species missing from these databases, we used other databases (mostly World Checklist of Selected Plant Families). For all species, author names and IDs for World Flora Online entries are provided to facilitate the alignment with other plant databases, and to avoid ambiguities. EpiList 1.0 will be a rich source for synthetic studies in ecology, biogeography, and evolutionary biology as it offers, for the first time, a species‐level overview over all currently known vascular epiphytes. At the same time, the list represents work in progress: species descriptions of epiphytic taxa are ongoing and published life form information in floristic inventories and trait and distribution databases is often incomplete and sometimes even wrong. Since the epiphytic growth blends into soil‐rooted growth and vice versa, the inclusion or exclusion of particular species in the current list will sometimes be contentious. Thus, initiating a well‐founded discussion was one of the motivations for compiling this database; our list represents 31,311 hypotheses on the life form of plant species, and we welcome feedback on possible omission or erroneous inclusions. We release these data into the public domain under a Creative Commons Zero license waiver. When you use the data in your publication, we request that you cite this data paper. If EpiList 1.0 is a major part of the data analyzed in your study, you may consider inviting the EpiList 1.0 core team as collaborators.

Vascular epiphytes are an important component of many ecosystems and constitute a substantial part of global plant diversity. In this context, accidental epiphytism, that is, the opportunistic epiphytic growth of typically terrestrial species, deserves special attention because it provides crucial insights into the global distribution of vascular epiphytes and the initial evolution of epiphytic lineages. Even though accidental epiphytes have been mentioned in the literature for more than a century, they have been neglected in most epiphyte studies. Only recently has accidental epiphytism been investigated more thoroughly. Therefore, the aim of this article is to provide a comprehensive review of the ecological basis and evolutionary relevance of this common but largely neglected phenomenon and to highlight open questions and promising research directions. Our central statement—that any species has the potential to grow epiphytically given the availability of suitable microhabitats and successful dispersal—is backed up by a compilation of observations of accidental epiphytes from numerous ecosystems with diverse climates, even including semiarid Mediterranean ones. A variety of arboreal microhabitats and environmental conditions conform to the ecological niche of typical terrestrial species, with the availability of such microhabitats depending on the interaction of local climate conditions, host tree age, and host species identity. Whenever suitable microhabitats are available in tree crowns, accidental epiphytism is limited primarily by dispersal. In an evolutionary context, the conquest of forest canopy represents an ecological opportunity where accidental epiphytes act as links between terrestrial and epiphytic life forms. We discuss two fundamental scenarios with sympatric speciation, selective pressure, autopolyploidy, and allopatric speciation as underlying mechanisms in the transition from terrestrial to epiphytic growth. In conclusion, we argue that accidental epiphytism is a substrate and dispersal-dependent phenomenon and that, both from an individual perspective and an evolutionary perspective, epiphytism reflects the occupation of suitable but previously unexploited arboreal microhabitats. Acknowledging the fundamental principles that plant growth is opportunistic and that dispersal is a stochastic process can decisively improve our understanding of species distributions and other ecological patterns, as in the case of accidental epiphytism.

The velamen radicum, a spongy, usually multiple epidermis of the roots, which at maturity consists of dead cells, is frequently described as an important adaptation of epiphytic orchids. Yet, quantitative evidence for the alleged functions, e.g., efficient water and nutrient uptake, nutrient retention, reduction of water loss, mechanical protection, or the avoidance of overheating, is rare or missing. We tested the notion originally put forward by Went in 1940 that the velamen allows plants to capture and immobilize the first solutions arriving in a rainfall, which are the most heavily charged with nutrients. In a series of experiments, we examined whether all necessary functional characteristics are given for this scenario to be realistic under ecological conditions. First, we show that the velamen of a large number of orchid species takes up solutions within seconds, while evaporation from the velamen takes several hours. Charged ions are retained in the velamen probably due to positive and negative charges in the cell walls, while uncharged compounds are lost to the external medium. Finally, we demonstrate that nutrient uptake follows biphasic kinetics with a highly efficient, active transport system at low external concentrations. Thus, our results lend strong support to Went's hypothesis: the velamen fulfills an important function in nutrient uptake in the epiphytic habitat. Most of the other functions outlined above still await similar experimental scrutiny.

Habitat heterogeneity is considered a primary causal driver underpinning patterns of diversity, yet the universal role of heterogeneity in structuring biodiversity is unclear due to a lack of coordinated experiments testing its effects across geographic scales and habitat types. Furthermore, key species interactions that can enhance heterogeneity, such as facilitation cascades of foundation species, have been largely overlooked in general biodiversity models. Here, we performed 22 geographically distributed experiments in different ecosystems and biogeographical regions to assess the extent to which variation in biodiversity is explained by three axes of habitat heterogeneity: the amount of habitat, its morphological complexity, and capacity to provide ecological resources (e.g. food) within and between co-occurring foundation species. We show that positive and additive effects across the three axes of heterogeneity are common, providing a compelling mechanistic insight into the universal importance of habitat heterogeneity in promoting biodiversity via cascades of facilitative interactions. Because many aspects of habitat heterogeneity can be controlled through restoration and management interventions, our findings are directly relevant to biodiversity conservation. Species interactions that can enhance habitat heterogeneity such as facilitation cascades of foundation species have been overlooked in biodiversity models. This study conducted 22 geographically distributed experiments in different ecosystems and biogeographical regions to assess the extent to which biodiversity is explained by three axes of habitat heterogeneity in facilitation cascades.

The extraordinary diversity of orchids has captivated scientists for more than a century, yet their complex spatial patterns at large scales remain poorly resolved. On islands, orchid diversity patterns are especially puzzling. While some islands are centres of orchid diversity, orchids are underrepresented on most islands. To disentangle such complex patterns, key functional differences among orchids must be considered – a distinction seldom made in biogeographical analyses. Using a global dataset of 454 islands, we tested prominent hypotheses in island biogeography, while simultaneously making the distinction between epiphytes and two terrestrial life forms (geophytes and non‐geophytes). Orchid diversity was unevenly distributed across islands and life forms. Epiphytic orchid diversity strongly increased with temperature, illustrating the near confinement of epiphytes to the tropics. Geophytes became proportionally more important with increasing seasonality, highlighting their ability to withstand harsh climatic conditions. Epiphytes and non‐geophytes both displayed responses (e.g. negative relationship with seasonality) related to their dependence on consistently favourable conditions, possibly because of the absence of subterranean storage organs. This highlights that the factors explaining orchid diversity differ strongly with, and are related to, life form. We suggest that key functional differences within and across plant families be considered in future studies to better understand drivers of complex diversity patterns.

The forest canopy is home to a rich biota. One salient feature are the dynamics of the habitat‐building trees, which are growing and eventually vanishing. Tree species strongly differ in growth rates, final size and longevity. Nevertheless, these inherent dynamics have been a blind spot in studies on host specificity of vascular epiphytes (vascular plants dwelling on trees without parasitizing them)—not least because tree growth rates and longevity are usually unknown in highly diverse tropical forests. The present study aims at tackling this blind spot. We compared epiphyte abundances (>23,000 individuals) found on 285 individuals of four focal tree species in a lowland moist forest in Panama. Data on repeated dbh censuses from a permanent tree plot provided the unique opportunity to estimate the age of our sampled trees. We compared the relative importance of tree longevity for host biases with that of other host tree characteristics, namely microclimatic conditions and bark acidity, rugosity and stability. The studied tree species differ in host quality and epiphyte species partly differ in host preferences. The conclusions concerning relative host tree quality depend hugely on whether or not different tree growth rates are considered. Comparing these conclusions allows important insights into the role of tree longevity in shaping epiphyte communities. Relating tree trait differences to the observed distributions of epiphytes among the focal tree species shows how the simultaneous action of various tree characteristics causes host biases. Synthesis. This study highlights the substantial but, up to now, hidden role of different tree growth rates for host tree specificity of vascular epiphytes. Future investigations need to consider this possibly confounding factor adequately to avoid spurious conclusions. Differences in tree growth rates have been a blind spot in studies on host specificity of vascular epiphytes. We compared epiphyte abundances on four tree species in a lowland moist forest. Host quality ranking depends hugely on whether tree size or age is used as a covariate. Future investigations need to consider different tree growth rates to avoid spurious conclusions.

In the field of biodiversity research, tree-related microhabitats in the temperate zone have received substantial interest in the last decade, but one particular microhabitat type, crown microsoils in tree crotches, has largely escaped the attention of scientists. We present a study from Central Europe that focused on the meso- and macrofauna in this microhabitat type. In twelve crotches we found more than 3000 individual animals out of 14 orders, encompassing all major elements of the terrestrial soil decomposer food web. Collembola accounted for more than 50% of the total. As expected, taxon richness correlated with habitat size. We conclude that this is an ideal natural system to study fundamental ecological questions of faunal community assembly.

Alpine treelines are expected to move upslope with a warming climate. However, so far treelines have responded inconsistently and future shifts remain difficult to predict since many factors unrelated to temperature, such as biotic interactions, affect responses at the local scale. Especially during the earliest regeneration stages, trees can be strongly influenced by alpine vegetation via both competition and facilitation. We aimed to understand the relative importance of these two types of interaction in different vegetation structures for treeline regeneration dynamics. Effects of herbaceous alpine vegetation on seedling emergence and first-year performance were studied in a field experiment in the French Alps (2100 m a.s.l.) with five important European treeline tree species: Larix decidua, Picea abies, Pinus cembra, Pinus uncinata and Sorbus aucuparia. Total emergence and locally-germinated seedling survival were not affected, but for seedlings planted at two months of age, negative vegetation impacts dominated for all response parameters: first-year survival, growth and carbohydrate accumulation. However, in the winter half-year, evergreen tree seedlings increased carbohydrate reserves under the protection of senescent herbs. Also, responses of locally-germinated seedlings suggest facilitative vegetation effects in the first two months after emergence. Thus, the interaction switched between competition and facilitation according to ontogenetic stage and seasons. Still, the net outcome after one year was negative, but species differed in their susceptibilities. Because initial establishment is the first bottleneck determining whether treelines remain stable or move upslope, understanding establishment, including site-, life-stage and species-specific processes, is essential for understanding observed treeline spatial patterns and dynamics. When developing predictive models of treeline dynamics, all these ‘local’ aspects should be incorporated in addition to more global drivers like changes in temperature.

Questions Vascular epiphytes make up about 9% of all vascular plants globally but are clearly underrepresented in the temperate zones. The accidental epiphytic occurrence of terrestrial species, in contrast, is common at these latitudes and can provide important insights in the evolution of obligate epiphytes. Here, we present the results of the first two annual censuses of a planned long‐term study on accidental epiphytes. We particularly aim to identify: (a) the abundance and species richness of accidental epiphytes; (b) the dynamics of accidental epiphytism; (c) occupied substrates and microsites; and (d) suitable host tree species. Location Harz Mountains, Germany, Central Europe. Methods We surveyed more than 1,200 trees in a low mountain range in two consecutive years for epiphytic individuals of vascular plants, considering host tree species and occupied microsites. Results About one quarter of the surveyed trees hosted epiphytic plants, totalling 1 450 (2016) and 1 350 (2017) individuals, respectively. These belonged to more than 100 species and 39 different families. The majority of epiphytic individuals newly emerged in 2017, whereas one‐third persisted since 2016 (or earlier) and a smaller proportion even reproduced in the epiphytic habitat. Accidental epiphytes were mostly restricted to host tree species providing water‐storing substrates such as extensive moss pads or arboreal soil accumulated in crotches. Compared to tropical epiphyte communities, accidental epiphytes in the temperate zones show a higher turnover, as they are less consistent in species abundance and composition in time. Conclusion For the majority of the observed species, epiphytism indeed is an accidental phenomenon. Undoubtedly, abiotic conditions limit the occurrence of obligate epiphytes at higher latitudes, but the presence and persistence of numerous epiphytic individuals illustrate that abiotic conditions do not per se preclude epiphytic occurrence of vascular plants in the north‐temperate zone. Besides water shortage, the availability of suitable host trees is a decisive environmental factor that contributes to limit epiphytism of vascular plants in Central Europe. True epiphytes are rare in the temperate zones; and accidental epiphytism of vascular plants, however, is common in higher latitudes and has been observed in more than 100 species in a Central European mountain range. Results indicate that, besides climatic conditions, the availability of suitable host trees contributes to limit epiphytism of vascular plants in the temperate zones.

Diatoms are of major importance in marine and freshwater systems, but their occurrence in terrestrial situations is generally thought to be exceptional. Following up on the accidental discovery of epiphytic diatoms on bark samples in an unrelated study, we investigated their presence in the tropical lowlands of Panama more systematically using scanning electron and light microscopy. We sampled inundated and aerial bark portions of Annona glabra, a tree that grows along the shore of Lake Gatun, and took bark samples from other tree and liana species at c. 1.5 m height in the forest understory. In total, we found 70 diatom taxa in 28 genera. Species numbers and composition differed among the three microhabitats with the largest numbers on inundated bark portions, but even in the forest understory, we found 12 taxa with densities of up to 900 frustules per mm[sup.−2] of bark. Our data set is still quite limited in scale but the results suggest the possibility that hitherto unacknowledged assemblages of epiphytic diatoms may be quite common in wet tropical forests, which clearly warrants further study.