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PREMISE Extrafloral nectaries are mainly studied in angiosperms, but have also been reported in 39 fern species. Here we provide a global review of nectaries in ferns, and study their structure, function, and nectar sugar composition in two genera. METHODS We searched in the literature and living plant collections of botanical gardens for indications of fern nectaries, studied the morpho-anatomy in the two genera Aglaomorpha and Campyloneurum, and analyzed the total sugar concentrations and ratios of 16 species. Diurnal nectar release was observed with time-lapse photography. RESULTS We found evidence for nectaries in 101 species of ferns from 11 genera and 6 families. Most of the nectary-bearing species were tree ferns (Cyatheaceae) and epiphytic ferns of the family Polypodiaceae. Nectaries consisted of cytoplasm-rich parenchyma with large nuclei and an epidermis with or without stomata, were attached to amphiphloic vascular bundles, and released nectar on the lower leaf surface mainly on expanding leaves during the night. Sugar concentrations varied between species (3.8-15.3%) but not between genera, and were sucrose-dominant (3 spp.), sucrose-rich (7), or hexose-rich (3). Under greenhouse conditions, introduced ants, scale insects, and snails fed on the nectar. CONCLUSIONS The wide taxonomic distribution, variable morphology, locations, and sugar compositions point to multiple evolutionary origins of fern nectaries. Nectar release in young leaves might attract mutualistic ants to protect leaves against herbivores only during this most vulnerable developmental stage. Even ex-situ, fern nectar is a valuable food source because it attracted several opportunistic animal species. This article is protected by copyright. All rights reserved.

The foliar surface forms one of the largest aboveground habitats on Earth and maintains plant-fungus relationships that greatly affect ecosystem functioning. Despite many studies with particular plant species, the foliar epiphytic mycobiome has not been studied across a large number of plant species from different taxa. Using high-throughput sequencing, we assessed epiphytic mycobiomes on leaf surfaces of 592 plant species in a botanical garden. Plants of angiosperms, gymnosperms, and pteridophytes were involved. Plant taxonomy, leaf side, growing environment, and evolutionary relationships were considered. We found that pteridophytes showed the higher fungal species diversity, stronger mutualistic fungal interactions, and a greater percentage of putative pathogens than gymnosperms and angiosperms. Plant taxonomic group, leaf side, and growing environment were significantly associated with the foliar epiphytic mycobiome, but the similarity of the mycobiomes among plants was not directly related to the distance of the host evolutionary tree. Our results provide a general understanding of the foliar fungal mycobiomes from pteridophytes to angiosperms. These findings will facilitate our understanding of foliar fungal epiphytes and their roles in plant communities and ecosystems.

International audience; Approximately 10% of vascular plants are epiphytes and, even though this has long been ignored in past research, are able to interact with a variety of fungi, including mycorrhizal taxa. However, the structure of fungal communities on bark, as well as their relationship with epiphytic plants, is largely unknown.To fill this gap, we conducted environmental metabarcoding of the ITS-2 region to understand the spatial structure of fungal communities of the bark of tropical trees, with a focus on epiphytic orchid mycorrhizal fungi, and tested the influence of root proximity.For all guilds, including orchid mycorrhizal fungi, fungal communities were more similar when spatially close on bark (i.e. they displayed positive spatial autocorrelation). They also showed distance decay of similarity with respect to epiphytic roots, meaning that their composition on bark increasingly differed, compared to roots, with distance from roots.We first showed that all of the investigated fungal guilds exhibited spatial structure at very small scales. This spatial structure was influenced by the roots of epiphytic plants, suggesting the existence of an epiphytic rhizosphere. Finally, we showed that orchid mycorrhizal fungi were aggregated around them, possibly as a result of reciprocal influence between the mycorrhizal partners.

The degeneration of submerged macrophytes and the invasion of Eichhornia crassipes (E. crassipes) destroyed the balance of aquatic ecosystems environments. In this study, responses of Vallisneria natans (V. natans) and the leaf-epiphytic biofilms to E. crassipes were analyzed to provide a technical scheme for V. natans restoration and E. crassipes control in eutrophic water. The results showed that a significant improvement of water quality achieved in 1100 ind·m-2 E. crassipes density group and TN removal rate reached 63.53%. The presence of E. crassipes changed the morphological characteristics of V. natans, which stimulated the adaptive mechanisms via promotion of shoot height and root length. Concentrations of the antioxidant enzymes, peroxidase, superoxide dismutase, and catalase in the V. natans leaves remained stable. But E. crassipes greatly increased the microbial diversity on V. natans leave biofilms. Furthermore, the greatest richness in bacterial community diversity was observed at 700, 1100, and 1200 ind·m-2 E. crassipes densities in heatmap, which was beneficial to the stability of the water ecological environment. These results showed that the combination of V. natans with E. crassipes of 1100 ind m-2 providing more favorable conditions for the growth and restoration of submerged macrophytes and improve the water quality. PRACTITIONER POINTS: The responses of submerged macrophytes to floating plants were studied. The optimal density of Eichhornia crassipes was 1100 ind m-2 . The biofilm microbial community changed in response to Eichhornia crassipes.

Bird‐mediated dispersal is presumed to be important in the dissemination of many different types of organisms, but concrete evidence remains scarce. This is especially true for biota producing microscopic propagules. Tree‐dwelling birds, such as woodpeckers, would seem to represent ideal dispersal vectors for organisms growing on standing tree trunks such as epiphytic lichens and fungi. Here, we utilize bird natural history collections as a novel source of data for studying dispersal ecology of plants, fungi, and microorganisms. We screened freshly preserved specimens of three Finnish woodpecker species for microscopic propagules. Samples were taken from bird feet, and chest and tail feathers. Propagules were extracted using a sonication–centrifugation protocol, and the material obtained was studied using light microscopy. Diverse biological material was recovered from all specimens of all bird species, from all positions sampled. Most abundant categories of discovered biological material included bryophyte fragments, fungal spores, and vegetative propagules of lichens. Also, freshwater diatoms, bryophyte spores, algal cells, testate amebae, rotifers, nematodes, pollen, and insect scales were identified. The method developed here is applicable to living specimens as well, making it a versatile tool for further research. Our findings highlight the potential of bird‐mediated dispersal for diverse organisms and showcase the use of natural history collections in ecological research., Woodpeckers are ideal candidates for bird‐mediated dispersal for a diverse set of organisms, especially epiphytes. We report high loads of diverse biological propagules extracted from feathers and feet of woodpecker specimens from natural history collections. The study highlights the potential of birds as dispersal vectors of lichens, fungi, bryophytes, and more.

Plant growth promotion has been found associated with plants on the surface (epiphytic), inside (endophytic), or close to the plant roots (rhizospheric). Endophytic bacteria mainly have been researched for their beneficial activities in terms of nutrient availability, plant growth hormones, and control of soil-borne and systemic pathogens. Molecular communications leading to these interactions between plants and endophytic bacteria are now being unrevealed using multidisciplinary approaches with advanced techniques such as metagenomics, metaproteomics, metatranscriptomics, metaproteogenomic, microRNAs, microarray, chips as well as the comparison of complete genome sequences. More than 400 genes in both the genomes of host plant and bacterial endophyte are up- or downregulated for the establishment of endophytism and plant growth-promoting activity. The involvement of more than 20 genes for endophytism, about 50 genes for direct plant growth promotion, about 25 genes for biocontrol activity, and about 10 genes for mitigation of different stresses has been identified in various bacterial endophytes. This review summarizes the progress that has been made in recent years by these modern techniques and approaches.

PREMISE Orchids depend primarily on mycorrhizal fungi to obtain nutrients throughout their life cycle. Epiphytic orchids account for 69% of orchid diversity. The unstable availability of water and nutrients in their arboreal habitats often results in severe water and nutrient stresses. Consequently, mycorrhizal associations may be important for the survival of epiphytic orchids, but our understanding thereof remains limited. Here, we investigated the mycorrhizal community in a single epiphytic orchid species, using more samples than in any previous study. METHODS We assessed the mycorrhizal communities of Thrixspermum japonicum, one of the most common epiphytic orchids in the temperate region of Japan. In total, 144 individuals were collected from 28 host tree species at 20 sites across 1300 km. The mycorrhizal fungi were identified based on nuclear ribosomal DNA internal transcribed spacer sequences and assigned operational taxonomic units (OTUs) based on 97% sequence similarity. RESULTS We obtained 24 OTUs; 9 belonged to the Ceratobasidiaceae and 15 to the Tulasnellaceae. These OTUs are widely distributed throughout the phylogenetic trees of the two fungal families. However, a single Ceratobasidiaceae OTU accounted for 49.7% of all fungal sequences and was predominant in samples from 15 host tree species and 12 sites. CONCLUSIONS Our results imply that despite having a broad range of mycorrhizal partners, T. japonicum was predominantly associated with a single fungal taxon at most of the sites among the host-tree species investigated. These findings contribute to elucidating mycorrhizal symbiosis in epiphytic habitats.

The regime of frequent small disturbances, such as the formation of tree‐fall gaps, is an important mechanism maintaining the diversity in tropical forests. Occurring primarily in forest gaps, or in riparian vegetation, ant gardens (hereafter AGs) are mutualistic associations involving several epiphytic plant species and ants. In the Amazon rainforest, a widespread AGs association is assembled by two aggressive species of parabiotic ants (Camponotus femoratus and Crematogaster levior). Working together, these ants aggressively monopolize food resources and attack any intruder on the AGs. Here, we demonstrated the extended protective effect of parabiotic ants on the leaf herbivory of neighboring plants, contrasting plants with and without extrafloral nectaries (hereafter EFNs). We performed the fieldwork in South Amazonia where we found 33 forest gaps, 16 with and 17 without AGs. Independently of the presence of AGs, plants possessing EFNs have less area removed by herbivores and fewer leaves with herbivore marks than those without EFNs. This decrease in herbivory was remarkable on EFN‐plants growing in gaps that also harbor AGs. Additional manipulative experiments confirmed this pattern: The AG‐ants were more efficient than no‐AG ants in removing live termite baits placed on EFN‐plants, but few attacks were observed on plants without EFNs. Thus, AG‐ants can extend their role in defending ant gardens from other plants growing in the same area, particularly those with EFNs. This extension of the mutualism benefits can affect the dynamics of the ecological succession and coevolutionary process in the Amazon forest. Abstract in Portuguese is available with online material

Concurrent anthropogenic global climate change and ocean acidification are expected to have a negative impact on calcifying marine organisms. While knowledge of biological responses of organisms to oceanic stress has emerged from single‐species experiments, these do not capture ecologically relevant scenarios where the potential for multi‐organism physiological interactions is assessed. Marine algae provide an interesting case study, as their photosynthetic activity elevates pH in the surrounding microenvironment, potentially buffering more acidic conditions for associated epiphytes. We present findings that indicate increased tolerance of an important epiphytic foraminifera, Marginopora vertebralis, to the effects of increased temperature (±3°C) and pCO2 (~1,000 µatm) when associated with its common algal host, Laurencia intricata. Specimens of M. vertebralis were incubated for 15 days in flow‐through aquaria simulating current and end‐of‐century temperature and pH conditions. Physiological measures of growth (change in wet weight), calcification (measured change in total alkalinity in closed bottles), photochemical efficiency (Fv/Fm), total chlorophyll, photosynthesis (oxygen flux), and respiration were determined. When incubated in isolation, M. vertebralis exhibited reduced growth in end‐of‐century projections of ocean acidification conditions, while calcification rates were lowest in the high‐temperature, low‐pH treatment. Interestingly, association with L. intricata ameliorated these stress effects with the growth and calcification rates of M. vertebralis being similar to those observed in ambient conditions. Total chlorophyll levels in M. vertebralis decreased when in association with L. intricata, while maximum photochemical efficiency increased in ambient conditions. Net production estimates remained similar between M. vertebralis in isolation and in association with L. intricata, although both production and respiration rates of M. vertebralis were significantly higher when associated with L. intricata. These results indicate that the association with L. intricata increases the resilience of M. vertebralis to climate change stress, providing one of the first examples of physiological buffering by a marine alga that can ameliorate the negative effects of changing ocean conditions., We present findings that indicate increased tolerance of an important epiphytic foraminifera, Marginopora vertebralis, to the antagonistic effects of increased temperature and pCO2, when associated with its common algal host, Laurencia intricata. When incubated in isolation, M. vertebralis exhibited reduced growth in end‐of‐century projections of ocean acidification conditions, while calcification rates were lowest in the high‐temperature, low‐pH treatment. Interestingly, association with L. intricata ameliorated these stress effects with the growth and calcification rates of M. vertebralis being similar to those observed in ambient conditions.

The expansion of angiosperm-dominated forests in the Cretaceous and early Cenozoic had a profound effect on terrestrial biota by creating novel ecological niches. The majority of modern fern lineages are hypothesized to have arisen in response to this expansion, particularly fern epiphytes that radiated into the canopy. Recent evidence, however, suggests that epiphytism does not correlate with increased diversification rates in ferns, calling into question the role of the canopy habitat in fern evolution. To understand the role of the canopy in structuring fern community diversity, we investigated functional traits of fern sporophytes and gametophytes across a broad phylogenetic sampling on the island of Moorea, French Polynesia, including > 120 species and representatives of multiple epiphytic radiations. While epiphytes showed convergence in small size and a higher frequency of noncordate gametophytes, they showed greater functional diversity at the community level relative to terrestrial ferns. These results suggest previously overlooked functional diversity among fern epiphytes, and raise the hypothesis that while the angiosperm canopy acted as a complex filter that restricted plant size, it also facilitated diversification into finely partitioned niches. Characterizing these niche axes and adaptations of epiphytic ferns occupying them should be a priority for future pteridological research.

Characterizing patterns in the microbial communities associated with canopy‐forming kelps is critical to understanding the mechanisms shaping macroalgal microbiomes. Using 16S rRNA gene sequencing, we characterized bacterial and archaeal communities associated with giant kelp in the Santa Barbara Channel. Our results indicate that kelp‐associated microbial communities are altered in the presence of epiphytic bryozoans and reflect changes previously observed in other studies of stressed macroalgae. This observation is indicative of patterns of microbiome disruption (dysbiosis) and opportunistic pathogenesis that may have implications for the health and productivity of foundational species of macroalgae. Our study highlights new patterns in host–microbiome associations and reveals the importance of considering host source and condition when investigating the dynamics of macroalgal microbiomes.

International audience; Little is known about direct interactions between diatoms. In this study, small diatoms attaching onto larger benthic and motile diatoms were considered. This phenomenon is commonly considered as epiphytism (with diatom epiphytes) although the exact nature of this interaction has not been investigated yet. The question that was addressed here is the specificity of the interaction, simply by observing species associations in their natural environment. A comparison of this special form of epiphytism was carried out between two sites, on a Mediterranean river and one of its tributaries. The results showed both host and epiphyte selection, but with some variability in species associations. This excludes the two alternative hypotheses that (i) large diatoms are a neutral substrate for smaller diatoms and that (ii) the interactions are strictly specific between two species.

The popular dual definition of lichen symbiosis is under question with recent findings of additional microbial partners living within the lichen body. Here we compare the distribution and co-occurrence patterns of lichen photobiont and recently described secondary fungus (Cyphobasidiales yeast) to evaluate their dependency on lichen host fungus (mycobiont). We sequenced the nuclear internal transcribed spacer (ITS) strands for mycobiont, photobiont, and yeast from six widespread northern hemisphere epiphytic lichen species collected from 25 sites in Switzerland and Estonia. Interaction network analyses and multivariate analyses were conducted on operational taxonomic units based on ITS sequence data. Our study demonstrates the frequent presence of cystobasidiomycete yeasts in studied lichens and shows that they are much less mycobiont-specific than the photobionts. Individuals of different lichen species growing on the same tree trunk consistently hosted the same or closely related mycobiont-specific Trebouxia lineage over geographic distances while the cystobasidiomycete yeasts were unevenly distributed over the study area - contrasting communities were found between Estonia and Switzerland. These results contradict previous findings of high mycobiont species specificity of Cyphobasidiales yeast at large geographic scales. Our results suggest that the yeast might not be as intimately associated with the symbiosis as is the photobiont.

Bird's nest ferns (Asplenium spp.) support large numbers of invertebrates, including centipedes. As top invertebrate predators, centipedes drive ecosystem function, for example, by regulating decomposer populations, but we know little of their ecology in forest canopies. We provide the first detailed observations of the diversity and structure of the centipede communities of bird's nest ferns, revealing the importance of these epiphytes as nurseries for centipedes. We collected 305 centipedes equating to ˜11,300 mg of centipede biomass from 44 bird's nest ferns (22 of which were from the high canopy and 22 from the low canopy) in primary tropical rainforest in Sabah, Malaysian Borneo. Most abundant were the Scolopendromorpha (n = 227 individuals), followed by the Geophilomorpha (n = 59), Lithobiomorpha (n = 14), and Scutigeromorpha (n = 5). Although we observed very little overlap in species between the forest strata, scolopendromorph centipedes dominated throughout the canopy. Null model analysis revealed no significant competitive interactions; on the contrary, we observed centipedes sharing nest sites within the ferns on three of the ten occasions that we found nests. All nests belonged to centipedes of the family Scolopendridae, which are typically aggressive, and usually show negative spatial association. This study reveals a diverse community of canopy centipedes, providing further evidence of the importance of bird's nest ferns to a wide range of animals, many of which use the ferns at critical life stages. Future conservation strategies should regard these ubiquitous epiphytes as umbrella species and protect them accordingly in landscape management decisions.

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

The filamentous red algal genus Gayliella, typified by G. flaccida, was segregated from Ceramium for a group of epiphytic species on the basis of morphological data and molecular phylogenetic analyses. However, while the genus was well investigated in some geographical areas, the treatment of the Brazilian taxa "Ceramium flaccidum" and "C. dawsonii" was incomplete and they were not transferred to Gayliella due to the problematic status of the specific epithets. Brazilian "C. flaccidum" did not fit within the circumscription of Gayliella flaccida; Brazilian "C. dawsonii" was positioned in three distinct lineages, none of which corresponded to known Gayliella species. Since the establishment of Gayliella, there has been no re-evaluation of Brazilian species, so this study revises the genus in Brazil using molecular and morphological data. Brazilian "C. flaccidum", "C. dawsonii", and other Brazilian collections were investigated by an integrative approach using phylogenetic analysis, genetic divergence, two species delimitation methods using three molecular markers (rbcL, cox1, and partial LSU), and comparative morphological and quantitative analyses. This molecular-assisted alpha taxonomy (MAAT) analysis placed Brazilian samples in five distinct well-supported lineages. For these, we propose a new combination, Gayliella dawsonii comb. nov. (lineage 2), based on analysis of type specimens and molecular data obtained from specimens collected near the type locality, and four new species: Gayliella ardissonei sp. nov. ("Ceramium flaccidum" from Brazil), G. iemanja sp. nov. ("C. dawsonii" lineage 3), G. tamoiensis sp. nov., and G. jolyana sp. nov. ("C. dawsonii" lineage 1).

The 18 O signature of atmospheric water vapour (δ18 OV ) is known to be transferred via leaf water to assimilates. It remains, however, unclear how the 18 O-signal transfer differs among plant species and growth forms. We performed a 9-hr greenhouse fog experiment (relative humidity ≥ 98%) with 18 O-depleted water vapour (-106.7‰) on 140 plant species of eight different growth forms during daytime. We quantified the 18 O-signal transfer by calculating the mean residence time of O in leaf water (MRTLW ) and sugars (MRTSugars ) and related it to leaf traits and physiological drivers. MRTLW increased with leaf succulence and thickness, varying between 1.4 and 10.8 hr. MRTSugars was shorter in C3 and C4 plants than in crassulacean acid metabolism (CAM) plants and highly variable among species and growth forms; MRTSugars was shortest for grasses and aquatic plants, intermediate for broadleaf trees, shrubs, and herbs, and longest for conifers, epiphytes, and succulents. Sucrose was more sensitive to δ18 OV variations than other assimilates. Our comprehensive study shows that plant species and growth forms vary strongly in their sensitivity to δ18 OV variations, which is important for the interpretation of δ18 O values in plant organic material and compounds and thus for the reconstruction of climatic conditions and plant functional responses.

PREMISE Conversion of primary forests to pastures is a major cause of habitat fragmentation in the tropics. Fragmentation is expected to impede gene flow for many plant species that are restricted to remaining forest fragments. Epiphytes may be especially vulnerable to this effect of forest fragmentation because they depend on host trees. However, trees that remain in pastures may enhance connectivity across the landscape for epiphyte species that can thrive on such trees. To investigate this possibility, we studied the genetic structures of two such species on isolated pasture trees and surrounding forest, in relation to their local abundances in different habitat types and aspects of their reproductive biology including pollen and seed dispersal agents, and looked for evidence of increased or diminished gene flow. METHODS We used microsatellite markers to assess geographic patterns of genetic diversity and differentiation in two epiphytic bromeliads, Catopsis nitida and Werauhia tonduziana, in the Monteverde region of Costa Rica. RESULTS About 85% of the FST value for Catopsis nitida was found among pastures within regions, while for Weruahia tonduziana, about 80% of the FST value was contributed by differences between regions, indicating much more gene flow within regions, relative to C. nitida. CONCLUSIONS Although there was substantial genetic differentiation among epiphyte populations, those on isolated pasture trees were not substantially less diverse than those in adjacent forests, suggesting that pasture trees may serve as "stepping stones" that help these species maintain their genetic connectedness and diversity at larger geographic scales.

Most plant communities consist of different structural and ecological subsets, ranging from cryptogams to different tree layers. The completeness and approach with which these subsets are sampled have implications for vegetation classification. Non‐vascular plants are often omitted or sometimes treated separately, referring to their assemblages as “synusiae” (e.g. epiphytes on bark, saxicolous species on rocks). The distinction of complete plant communities (phytocoenoses or holocoenoses) from their parts (synusiae or merocoenoses) is crucial to avoid logical problems and inconsistencies of the resulting classification systems. We here describe theoretical differences between the phytocoenosis as a whole and its parts, and outline consequences of this distinction for practise and terminology in vegetation classification. To implement a clearer separation, we call for modifications of the International Code of Phytosociological Nomenclature and the EuroVegChecklist. We believe that these steps will make vegetation classification systems better applicable and raise the recognition of the importance of non‐vascular plants in the vegetation as well as their interplay with vascular plants.

Few studies have explored how - within the same system - clonality and positive plant-plant interactions might interact to regulate plant community composition. Canopy-dwelling epiphytes in species-rich forests provide an ideal system for studying this because many epiphytic vascular plants undertake clonal growth and because vascular epiphytes colonize canopy habitats after the formation of nonvascular epiphyte (i.e. bryophyte and lichen) mats. We investigated how clonal integration of seven dominant vascular epiphytes influenced inter-specific interactions between vascular epiphytes and nonvascular epiphytes in a subtropical montane moist forest in southwest China. Both clonal integration and environmental buffering from nonvascular epiphytes increased survival and growth of vascular epiphytes. The benefits of clonal integration for vascular epiphytes were higher when nonvascular epiphytes were removed. Similarly, facilitation from nonvascular epiphytes played a more important role when clonal integration of vascular epiphytes was eliminated. Overall, clonal integration had greater benefits than inter-specific facilitation. This study provides novel evidence for interactive effects of clonality and facilitation between vascular and nonvascular species, and has implications for our understanding of a wide range of ecosystems where both high levels of clonality and facilitation are expected to occur.

Decomposition is a major component of global carbon cycling. However, approximately 50% of wood necromass and a small proportion of leaf litter do not contact the forest floor, and the factors that regulate the decomposition above the forest floor are largely untested. We hypothesized that separation from soil resources causes slower decomposition rates above the forest floor. Specifically, we tested whether slower decomposition results from decreased nutrient availability (the nutrient limitation hypothesis) and/or microbial dispersal limitation (the dispersal limitation hypothesis) in the absence of soil resources. We tested these hypotheses by combining experimental manipulations of epiphytes and macronutrient fertilization with elemental analyses and community metabarcoding (fungi and prokaryotes). Specifically, we compared wood stick and cellulose decomposition among three treatments: an unaltered trunk section, an epiphyte mat, and a ‘removal treatment’ where an epiphyte mat was removed to test the effect of soil resources. We also performed a factorial fertilization experiment to test the effects of nitrogen (N) and phosphorus (P) on the decomposition of suspended cellulose. Decomposition rates were fastest on the epiphyte mats, intermediate in the removal treatment and slowest in the controls. Phosphorus addition increased decomposition rates in the fertilization experiment, and greater P concentrations, along with some micronutrients, were associated with increased rates of decomposition on the epiphyte mats and in the removal treatments. Locally dispersed fungi dominated the wood stick communities, indicating that fungal dispersal is limited in the canopy, and fungal saprotrophs were associated with increased rates of decomposition on the epiphytes. These experiments show that slowed decomposition above the forest floor is caused, in part, by separation from soil resources. Moreover, our findings provide support for both the nutrient limitation and dispersal limitation hypotheses and indicate that mechanisms regulating canopy‐level decomposition differ from those documented on the forest floor. This demonstrates the need for a holistic approach to decomposition that considers the vertical position of necromass as it decomposes. Further experimentation is necessary to quantify interactions between community assembly processes, nutrient availability, substrate traits, and microclimate. A free Plain Language Summary can be found within the Supporting Information of this article.

Coastal eutrophication is a growing problem worldwide, leading to increased epiphyte overgrowth of seagrass leaves. Yet little is known about how epiphytes affect key biogeochemical conditions and processes in the seagrass phyllosphere. We used electrochemical microsensors to measure microgradients of O2, pH, and CO2 at the bare and epiphyte-covered leaf surface of seagrass (Zostera marina L.) to determine effects of epiphytes on the leaf chemical microenvironment. Epiphytes result in extreme daily fluctuations in pH, O2, and inorganic carbon concentrations at the seagrass leaf surface severely hampering the plant's performance. In light, leaf epiphyte biofilms and their diffusive boundary layer lead to strong basification, markedly reducing the CO2 and HCO3 - availability at the leaf surface, leading to reduced photosynthetic efficiency as a result of carbon limitation and enhanced photorespiration. With epiphytes, leaf surface pH increased to >10, thereby exceeding final pH levels (~9.62) and CO2 compensation points for active photosynthesis. In darkness, epiphyte biofilms resulted in increased CO2 and hypoxia at the leaf surface. Epiphytes can lead to severe carbon limitation in seagrasses owing to strong phyllosphere basification leading to CO2 depletion and costly, yet limiting, HCO3 - utilization, increasing the risk of plant starvation.

QUESTION: Biodiversity surveys of epiphytic bryophytes and lichens on standing trees are mostly restricted to the lowermost 2 m, since sampling above this level is time‐consuming and therefore expensive. However, sampling only the stem base is likely to result in underestimation of forest epiphyte diversity, because microclimate and physico‐chemical site conditions, both of which vary with height above the ground, play important roles for poikilohydric organisms. We investigate variation in epiphyte diversity and composition along the vertical gradient in forests and discuss factors that may potentially account for height‐dependent distribution patterns. LOCATION: Montane primeval beech forest Havesova in the Carpathians, eastern Slovakia. METHODS: Ten circular plots, each 500 m², were selected randomly in an area severely affected by windfall. Epiphytes were sampled along the stems of recently uprooted trees, from base to the canopy. Mean Ellenberg indicator values (EIV) for light, moisture, acidity and nitrogen availability were used as surrogate variables for environmental interpretation of compositional patterns for sampling segments of 2 m along the stem. RESULTS: By sampling only the lowest 2 m, 10% of the total bryophyte and 48% of the lichen species would have been missed. We detected systematic patterns of compositional variation for both groups along the vertical gradient. While pleurocarpous mosses and liverworts were mostly restricted to the stem base, acrocarpous mosses and crustose lichens occurred along the stem almost to the top. Foliose and fruticose lichens were concentrated at intermediate heights and in the upper canopy. Patterns of variation in EIVs suggested that increasing light availability, decreasing moisture and increasing inputs of acidic and nitrous substances are responsible for the compositional shifts along the vertical gradient. CONCLUSIONS: Sampling only the stem base implies severe underestimation of the epiphyte diversity in temperate broad‐leaved forests. The middle and the upper parts of tree stems have distinctive compositions of epiphytic cryptogams, because microclimate and physico‐chemical site factors vary with height above the ground.

Premise Because of its broad range in the neotropical rainforest and within tree canopies, the tank bromeliad Guzmania monostachia was investigated as a model of how varying leaf hydraulic conductance (Kleaf ) could help plants resist and recover from episodic drought. The two pathways of Kleaf , inside and outside the xylem, were also examined to determine the sites and causes of major hydraulic resistances within the leaf. Methods We measured leaf hydraulic conductance for plants in the field and laboratory under wet, dry, and rewetted conditions and applied physiological, anatomical, and gene expression analysis with modeling to investigate changes in Kleaf . Results After 7 d with no rain in the field or 14 days with no water in the glasshouse, Kleaf decreased by 50% yet increased to hydrated values within 4 d of tank refilling. Staining to detect embolism combined with modeling indicated that changes outside the xylem were of greater importance to Kleaf than were changes inside the xylem and were associated with changes in intercellular air spaces (aerenchyma), aquaporin expression and inhibition, and cuticular conductance. Conclusions Low values for all conductances during drying, particularly in pathways outside the xylem, lead to hydraulic resilience for this species and may also contribute to its broad environmental tolerances.

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.