Adaptive evolution of the enigmatic Takakia now facing climate change in Tibet.

The most extreme environments are the most vulnerable to transformation under a rapidly changing climate. These ecosystems harbor some of the most specialized species, which will likely suffer the highest extinction rates. We document the steepest temperature increase (2010-2021) on record at altitudes of above 4,000 m, triggering a decline of the relictual and highly adapted moss Takakia lepidozioides. Its de-novo-sequenced genome with 27,467 protein-coding genes includes distinct adaptations to abiotic stresses and comprises the largest number of fast-evolving genes under positive selection. The uplift of the study site in the last 65 million years has resulted in life-threatening UV-B radiation and drastically reduced temperatures, and we detected several of the molecular adaptations of Takakia to these environmental changes. Surprisingly, specific morphological features likely occurred earlier than 165 mya in much warmer environments. Following nearly 400 million years of evolution and resilience, this species is now facing extinction.

Plasmodesmata dynamics in bryophyte model organisms: secondary formation and developmental modifications of structure and function.

MAIN CONCLUSION: Developing bryophytes differentially modify their plasmodesmata structure and function. Secondary plasmodesmata formation via twinning appears to be an ancestral trait. Plasmodesmata networks in hornwort sporophyte meristems resemble those of angiosperms. All land-plant taxa use plasmodesmata (PD) cell connections for symplasmic communication. In angiosperm development, PD networks undergo an extensive remodeling by structural and functional PD modifications, and by postcytokinetic formation of additional secondary PD (secPD). Since comparable information on PD dynamics is scarce for the embryophyte sister groups, we investigated maturating tissues of Anthoceros agrestis (hornwort), Physcomitrium patens (moss), and Marchantia polymorpha (liverwort). As in angiosperms, quantitative electron microscopy revealed secPD formation via twinning in gametophytes of all model bryophytes, which gives rise to laterally adjacent PD pairs or to complex branched PD. This finding suggests that PD twinning is an ancient evolutionary mechanism to adjust PD numbers during wall expansion. Moreover, all bryophyte gametophytes modify their existing PD via taxon-specific strategies resembling those of angiosperms. Development of type II-like PD morphotypes with enlarged diameters or formation of pit pairs might be required to maintain PD transport rates during wall thickening. Similar to angiosperm leaves, fluorescence redistribution after photobleaching revealed a considerable reduction of the PD permeability in maturating P. patens phyllids. In contrast to previous reports on monoplex meristems of bryophyte gametophytes with single initials, we observed targeted secPD formation in the multi-initial basal meristems of A. agrestis sporophytes. Their PD networks share typical features of multi-initial angiosperm meristems, which may hint at a putative homologous origin. We also discuss that monoplex and multi-initial meristems may require distinct types of PD networks, with or without secPD formation, to control maintenance of initial identity and positional signaling.

Impact of changing climate on bryophyte contributions to terrestrial water, carbon, and nitrogen cycles.

Bryophytes, including the lineages of mosses, liverworts, and hornworts, are the second-largest photoautotroph group on Earth. Recent work across terrestrial ecosystems has highlighted how bryophytes retain and control water, fix substantial amounts of carbon (C), and contribute to nitrogen (N) cycles in forests (boreal, temperate, and tropical), tundra, peatlands, grasslands, and deserts. Understanding how changing climate affects bryophyte contributions to global cycles in different ecosystems is of primary importance. However, because of their small physical size, bryophytes have been largely ignored in research on water, C, and N cycles at global scales. Here, we review the literature on how bryophytes influence global biogeochemical cycles, and we highlight that while some aspects of global change represent critical tipping points for survival, bryophytes may also buffer many ecosystems from change due to their capacity for water, C, and N uptake and storage. However, as the thresholds of resistance of bryophytes to temperature and precipitation regime changes are mostly unknown, it is challenging to predict how long this buffering capacity will remain functional. Furthermore, as ecosystems shift their global distribution in response to changing climate, the size of different bryophyte-influenced biomes will change, resulting in shifts in the magnitude of bryophyte impacts on global ecosystem functions.

The bryophyte rhizoid-sphere microbiome responds to water deficit.

The roots of vascular plants are colonised by a multitude of microbes, which play an important role in plant health and stress resilience. Drought stress in particular is devastating for crop yield and causes major shifts in the rhizosphere microbial communities. However, the microbiome associated to the rhizoids (hereafter termed rhizoid-sphere) of the nonvascular bryophytes remains largely unexplored. Here, we use amplicon sequencing to explore the rhizoid-sphere microbiome of three bryophyte species under drought and well-watered conditions. Comparing rhizoid-sphere microbial communities associated with the two liverworts Marchantia polymorpha and Marchantia paleacea and the moss Physcomitrium patens showed characteristic differences in composition between host species and both conserved and unique changes under drought. At phylum level, these changes were similar to changes in the rhizosphere of angiosperms under drought. Furthermore, we observed strong differences in rhizoid-sphere colonisation between bryophyte species for taxa known for nitrogen fixation and plant growth promotion. Interestingly, M. polymorpha prioritised the growth of belowground organs under osmotic stress, as is the case for angiosperms under drought. Taken together, our results show interesting parallels between bryophytes and angiosperms in the relation with their rhizo(id-)sphere, suggesting evolutionary conservation among land plants in their response to drought stress.

Moss species and precipitation mediate experimental warming stimulation of growing season N2 fixation in subarctic tundra.

Climate change in high latitude regions leads to both higher temperatures and more precipitation but their combined effects on terrestrial ecosystem processes are poorly understood. In nitrogen (N) limited and often moss-dominated tundra and boreal ecosystems, moss-associated N2 fixation is an important process that provides new N. We tested whether high mean annual precipitation enhanced experimental warming effects on growing season N2 fixation in three common arctic-boreal moss species adapted to different moisture conditions and evaluated their N contribution to the landscape level. We measured in situ N2 fixation rates in Hylocomium splendens, Pleurozium schreberi and Sphagnum spp. from June to September in subarctic tundra in Sweden. We exposed mosses occurring along a natural precipitation gradient (mean annual precipitation: 571-1155 mm) to 8 years of experimental summer warming using open-top chambers before our measurements. We modelled species-specific seasonal N input to the ecosystem at the colony and landscape level. Higher mean annual precipitation clearly increased N2 fixation, especially during peak growing season and in feather mosses. For Sphagnum-associated N2 fixation, high mean annual precipitation reversed a small negative warming response. By contrast, in the dry-adapted feather moss species higher mean annual precipitation led to negative warming effects. Modelled total growing season N inputs for Sphagnum spp. colonies were two to three times that of feather mosses at an area basis. However, at the landscape level where feather mosses were more abundant, they contributed 50% more N than Sphagnum. The discrepancy between modelled estimates of species-specific N input via N2 fixation at the moss core versus ecosystem scale, exemplify how moss cover is essential for evaluating impact of altered N2 fixation. Importantly, combined effects of warming and higher mean annual precipitation may not lead to similar responses across moss species, which could affect moss fitness and their abilities to buffer environmental changes.

Cell death in bryophytes: emerging models to study core regulatory modules and conserved pathways.

This review summarizes recent progress in our current understanding of the mechanisms underlying the cell death pathways in bryophytes, focusing on conserved pathways and particularities in comparison to angiosperms. Regulated cell death (RCD) plays key roles during essential processes along the plant life cycle. It is part of specific developmental programmes and maintains homeostasis of the organism in response to unfavourable environments. Bryophytes could provide valuable models to study developmental RCD processes as well as those triggered by biotic and abiotic stresses. Some pathways analogous to those present in angiosperms occur in the gametophytic haploid generation of bryophytes, allowing direct genetic studies. In this review, we focus on such RCD programmes, identifying core conserved mechanisms and raising new key questions to analyse RCD from an evolutionary perspective.

Dehydration and rehydration differently affect photosynthesis and volatile monoterpenes in bryophytes with contrasting ecological traits.

Bryophytes desiccate rapidly when relative humidity decreases. The capacity to withstand dehydration depends on several ecological and physiological factors. Volatile organic compounds (VOCs) may have a role in enhancing tolerance to desiccating bryophytes. However, the functions of VOCs in bryophytes have received little attention so far. We aimed to investigate the impact of a dehydration-rehydration treatment on primary carbon metabolism and volatile terpenes (VTs) in three bryophytes with contrasting ecological traits: Vessicularia dubyana, Porella platyphylla and Pleurochaete squarrosa. First, we confirmed the desiccation sensitivity gradient of the species. Under fully hydrated conditions, the photosynthetic rate (A) was inversely associated with stress tolerance, with a lower rate in more tolerant species. The partial recovery of A in P. platyphylla and P. squarrosa after rehydration confirmed the desiccation tolerance of these two species. On the other hand, A did not recover after rehydration in V. dubyana. Regarding VT, each species exhibited a distinct VT profile under optimum hydration, with the highest VT pool found in the more desiccation-sensitive species (V. dubyana). However, the observed species-specific VT pattern could be associated with the ecological habitat of each species. P. squarrosa, a moss of dry habitats, may synthesize mainly non-volatile secondary metabolites as stress-defensive compounds. On the other hand, V. dubyana, commonly found submerged, may need to invest photosynthetically assimilated carbon to synthesize a higher amount of VTs to cope with transient water stress occurrence. Further research on the functions of VTs in bryophytes is needed to deepen our understanding of their ecological significance.

Terrestrial mosses as a substrate and potential host for cyanobacteria, green algae and diatoms.

Most studies of terrestrial bryophytes as natural substrates for photosynthetic microorganisms have been performed in the polar regions, where bryophytes are an important part of the ecosystem. As they remain green throughout the year, bryophytes may also be an ideal substrate for epiphytic organisms in temperate regions. The present study investigated the colonization potential and diversity of microalgae on selected plant species in riparian forest and spruce monoculture in a temperate region. It examines whether the presence of algae is related to substrate humidity, the micromorphology of gametophyte or the seasonal availability of substrate. The taxonomic diversity of algae was studied. Cyanobacteria and green algae were cultured on BG-11 agar medium, while diatoms were identified in permanent diatomaceous slides. The alpha- and beta-diversity indices were calculated, and the communities were compared using Bray-Curtis distances and multidimensional correspondence analyses. Our findings indicate that the largest number of alga species were diatoms; however, their presence was only observed in riparian forest and was associated with high humidity. Both aerophilic and freshwater taxa were noted, the latter carried by water from nearby aquatic ecosystem. Green algae were present in both phytocoenoses and humidity appears to have no substantial effect on the degree of colonization; their diversity was low and the group consisted of terrestrial taxa. In two bryophytes growing at the highest humidity, cyanobacteria were only identified in culture. The key factor influencing the degree of microalgae colonization was the humidity of the substrate, which was related to the distance from water.

A research agenda for nonvascular photoautotrophs under climate change.

Nonvascular photoautotrophs (NVP), including bryophytes, lichens, terrestrial algae, and cyanobacteria, are increasingly recognized as being essential to ecosystem functioning in many regions of the world. Current research suggests that climate change may pose a substantial threat to NVP, but the extent to which this will affect the associated ecosystem functions and services is highly uncertain. Here, we propose a research agenda to address this urgent question, focusing on physiological and ecological processes that link NVP to ecosystem functions while also taking into account the substantial taxonomic diversity across multiple ecosystem types. Accordingly, we developed a new categorization scheme, based on microclimatic gradients, which simplifies the high physiological and morphological diversity of NVP and world-wide distribution with respect to several broad habitat types. We found that habitat-specific ecosystem functions of NVP will likely be substantially affected by climate change, and more quantitative process understanding is required on: (1) potential for acclimation; (2) response to elevated CO2 ; (3) role of the microbiome; and (4) feedback to (micro)climate. We suggest an integrative approach of innovative, multimethod laboratory and field experiments and ecophysiological modelling, for which sustained scientific collaboration on NVP research will be essential.

Disentangling climate change from air pollution effects on epiphytic bryophytes.

At the interface between atmosphere and vegetation, epiphytic floras have been largely used as indicators of air quality. The recovery of epiphytes from high levels of SO2 pollution has resulted in major range changes, whose interpretation has, however, been challenged by concomitant variation in other pollutants as well as climate change. Here, we combine historical and contemporary information on epiphytic bryophyte species distributions, climatic conditions, and pollution loads since the 1980s in southern Belgium to disentangle the relative impact of climate change and air pollution on temporal shifts in species composition. The relationship between the temporal variation of species composition, climatic conditions, SO2 , NO2 , O3 , and fine particle concentrations, was analyzed by variation partitioning. The temporal shift in species composition was such, that it was, on average, more than twice larger than the change in species composition observed today among communities scattered across the study area. The main driver, contributing to 38% of this temporal shift in species composition, was the variation of air quality. Climate change alone did not contribute to the substantial compositional shifts in epiphytic bryophyte communities in the course of the last 40 years. As a consequence of the substantial drop of N and S loads over the last decades, present-day variations of epiphytic floras were, however, better explained by the spatial variation of climatic conditions than by extant pollution loads. The lack of any signature of recolonization delays of formerly polluted areas in the composition of modern floras suggests that epiphytic bryophytes efficiently disperse at the landscape scale. We suggest that a monitoring of epiphyte communities at 10-year intervals would be desirable to assess the impact of raising pollution sources, and especially pesticides, whose impact on bryophytes remains poorly documented.

Chemical Stimulation of Heterocyte Differentiation by the Feather Moss Hylocomium splendens: a Potential New Step in Plant-Cyanobacteria Symbioses.

Cyanobacteria associated with mosses play a key role in the nitrogen (N) cycle in unpolluted ecosystems. Mosses have been found to release molecules that induce morphophysiological changes in epiphytic cyanobionts. Nevertheless, the extent of moss influence on these microorganisms remains unknown. To evaluate how mosses or their metabolites influence N2 fixation rates by cyanobacteria, we assessed the nitrogenase activity, heterocyte frequency and biomass of a cyanobacterial strain isolated from the feather moss Hylocomium splendens and a non-symbiotic strain when they were either growing by themselves, together with H. splendens or exposed to H. splendens water, acetone, ethanol, or isopropanol extracts. The same cyanobacterial strains were added to another moss (Taxiphyllum barbieri) and a liverwort (Monosolenium tenerum) to assess if these bryophytes affect N2 fixation differently. Although no significant increases in nitrogenase activity by the cyanobacteria were observed when in contact with H. splendens shoots, both the symbiotic and non-symbiotic cyanobacteria increased nitrogenase activity as well as heterocyte frequency significantly upon exposure to H. splendens ethanol extracts. Contact with T. barbieri shoots, on the other hand, did lead to increases in nitrogenase activity, indicating low host-specificity to cyanobacterial activity. These findings suggest that H. splendens produces heterocyte-differentiating factors (HDFs) that are capable of stimulating cyanobacterial N2 fixation regardless of symbiotic competency. Based on previous knowledge about the chemical ecology and dynamics of moss-cyanobacteria interactions, we speculate that HDF expression by the host takes place in a hypothetical new step occurring after plant colonization and the repression of hormogonia.

Sub-arctic mosses and lichens show idiosyncratic responses to combinations of winter heatwaves, freezing and nitrogen deposition.

Arctic ecosystems are increasingly exposed to extreme climatic events throughout the year, which can affect species performance. Cryptogams (bryophytes and lichens) provide important ecosystem services in polar ecosystems but may be physiologically affected or killed by extreme events. Through field and laboratory manipulations, we compared physiological responses of seven dominant sub-Arctic cryptogams (three bryophytes, four lichens) to single events and factorial combinations of mid-winter heatwave (6°C for 7 days), re-freezing, snow removal and summer nitrogen addition. We aimed to identify which mosses and lichens are vulnerable to these abiotic extremes and if combinations would exacerbate physiological responses. Combinations of extremes resulted in stronger species responses but included idiosyncratic species-specific responses. Species that remained dormant during winter (March), irrespective of extremes, showed little physiological response during summer (August). However, winter physiological activity, and response to winter extremes, was not consistently associated with summer physiological impacts. Winter extremes affect cryptogam physiology, but summer responses appear mild, and lichens affect the photobiont more than the mycobiont. Accounting for Arctic cryptogam response to multiple climatic extremes in ecosystem functioning and modelling will require a better understanding of their winter eco-physiology and repair capabilities.

Desiccation-rehydration measurements in bryophytes: current status and future insights.

Desiccation-rehydration experiments have been employed over the years to evaluate desiccation tolerance of bryophytes (Bryophyta, Marchantiophyta, and Anthocerotophyta). Researchers have applied a spectrum of protocols to induce desiccation and subsequent rehydration, and a wide variety of techniques have been used to study desiccation-dependent changes in bryophyte molecular, cellular, physiological, and structural traits, resulting in a multifaceted assortment of information that is challenging to synthesize. We analysed 337 desiccation-rehydration studies, providing information for 351 species, to identify the most frequent methods used, analyse the advances in desiccation studies over the years, and characterize the taxonomic representation of the species assessed. We observed certain similarities across methodologies, but the degree of convergence among the experimental protocols was surprisingly low. Out of 52 bryophyte orders, 40% have not been studied, and data are lacking for multiple remote or difficult to access locations. We conclude that for quantitative interspecific comparisons of desiccation tolerance, rigorous standardization of experimental protocols and measurement techniques, and simultaneous use of an array of experimental techniques are required for a mechanistic insight into the different traits modified in response to desiccation. New studies should also aim to fill gaps in taxonomic, ecological, and spatial coverage of bryophytes.

Ecology and responses to climate change of biocrust-forming mosses in drylands.

Interest in understanding the role of biocrusts as ecosystem engineers in drylands has substantially increased during the past two decades. Mosses are a major component of biocrusts and dominate their late successional stages. In general, their impacts on most ecosystem functions are greater than those of early-stage biocrust constituents. However, it is common to find contradictory results regarding how moss interactions with different biotic and abiotic factors affect ecosystem processes. This review aims to (i) describe the adaptations and environmental constraints of biocrust-forming mosses in drylands, (ii) identify their primary ecological roles in these ecosystems, and (iii) synthesize their responses to climate change. We emphasize the importance of interactions between specific functional traits of mosses (e.g. height, radiation reflectance, morphology, and shoot densities) and both the environment (e.g. climate, topography, and soil properties) and other organisms to understand their ecological roles and responses to climate change. We also highlight key areas that should be researched in the future to fill essential gaps in our understanding of the ecology and the responses to ongoing climate change of biocrust-forming mosses. These include a better understanding of intra- and interspecific interactions and mechanisms driving mosses' carbon balance during desiccation-rehydration cycles.

Biological soil crusts and how they might colonize other worlds: insights from these Brazilian ecosystem engineers.

When bryophytes, lichens, eukaryotic algae, cyanobacteria, bacteria, and fungi live interacting intimately with the most superficial particles of the soil, they form a complex community of organisms called the biological soil crust (BSC or biocrust). These biocrusts occur predominantly in drylands, where they provide important ecological services such as soil aggregation, moisture retention, and nitrogen fixation. Unfortunately, many BSC communities remain poorly explored, especially in the tropics. This review summarizes studies about BSCs in Brazil, a tropical megadiverse country, and shows the importance of ecological, physiological, and taxonomic knowledge of biocrusts. We also compare Brazilian BSC communities with others around the world, describe why BSCs can be considered ecosystem engineers, and propose their use in the colonization of other worlds.

Biocrust impacts on dryland soil water balance: A path toward the whole picture.

As a crucial living feature inhabiting the soil-atmosphere boundary, biocrusts play a vital role in liquid water or vapor transport through surface soil and thus have strong effects on soil water regimes. However, it remains unclear how biocrusts affect annual or multiyear soil water budgets through the regulation of evaporation outputs and non-rainfall water (NRW) or infiltration inputs. Thus, we used automated microlysimeters to continually investigate the differences in evaporation and NRW rates between moss-dominated biocrusts and bare soil at 0-5 cm depth for 2 years. The upper 30 cm of soil moisture (θ) and water storage (W) of bare soil and biocrusts were also monitored. Our results showed that the daily evaporation rate (E) of biocrusts was 17% higher than bare soil. Especially after rainfall events, biocrusts had higher E and larger cumulative evaporation than bare soil. Besides, the daily NRW of biocrusts averaged 15% higher than bare soil over 2 years. Furthermore, biocrusts increased θ by 11%-76% at 0-10 cm depth but decreased θ by 32%-56% at 20-30 cm depth in comparison to bare soil, and they subsequently decreased W by 20% at 0-30 cm depth. Summarized annually, the NRW amount of biocrusts was 19% higher than bare soil, but at the same time, the cumulative evaporation of biocrusts was also 19% higher than bare soil. Finally, biocrusts resulted in more water loss at shallow depth through evaporation and lessened total W throughout 0-30 cm depth of soil. These findings demonstrate that although biocrusts input more NRW into surface soil, these water inputs partially offset their intensified evaporation. Given that all rainfall water infiltrates into the soil in our study system, our findings indicate that biocrusts may have an overall negative effect on soil water balance there, while at the same time increasing water storage and availability of the deeper soil underlying biocrusts.

Photosynthesis and photosynthetic efficiencies along the terrestrial plant's phylogeny: lessons for improving crop photosynthesis.

Photosynthesis is the basis of all life on Earth. Surprisingly, until very recently, data on photosynthesis, photosynthetic efficiencies, and photosynthesis limitations in terrestrial land plants other than spermatophytes were very scarce. Here we provide an updated data compilation showing that maximum photosynthesis rates (expressed either on an area or dry mass basis) progressively scale along the land plant's phylogeny, from lowest values in bryophytes to largest in angiosperms. Unexpectedly, both photosynthetic water (WUE) and nitrogen (PNUE) use efficiencies also scale positively through the phylogeny, for which it has been commonly reported that these two efficiencies tend to trade-off between them when comparing different genotypes or a single species subject to different environmental conditions. After providing experimental evidence that these observed trends are mostly due to an increased mesophyll conductance to CO2 - associated with specific anatomical changes - along the phylogeny, we discuss how these findings on a large phylogenetic scale can provide useful information to address potential photosynthetic improvements in crops in the near future.

Photosynthesis on the edge: photoinhibition, desiccation and freezing tolerance of Antarctic bryophytes.

In Antarctica, multiple stresses (low temperatures, drought and excessive irradiance) hamper photosynthesis even in summer. We hypothesize that controlled inactivation of PSII reaction centres, a mechanism widely studied by pioneer work of Fred Chow and co-workers, may effectively guarantee functional photosynthesis under these conditions. Thus, we analysed the energy partitioning through photosystems in response to temperature in 15 bryophyte species presenting different worldwide distributions but all growing in Livingston Island, under controlled and field conditions. We additionally tested their tolerance to desiccation and freezing and compared those with their capability for sexual reproduction in Antarctica (as a proxy to overall fitness). Under field conditions, when irradiance rules air temperature by the warming of shoots (up to 20 °C under sunny days), a predominance of sustained photoinhibition beyond dynamic heat dissipation was observed at low temperatures. Antarctic endemic and polar species showed the largest increases of photoinhibition at low temperatures. On the contrary, the variation of thermal dissipation with temperature was not linked to species distribution. Instead, maximum non-photochemical quenching at 20 °C was related (strongly and positively) with desiccation tolerance, which also correlated with fertility in Antarctica, but not with freezing tolerance. Although all the analysed species tolerated - 20 °C when dry, the tolerance to freezing in hydrated state ranged from the exceptional ability of Schistidium rivulare (that survived for 14 months at - 80 °C) to the susceptibility of Bryum pseudotriquetrum (that died after 1 day at - 20 °C unless being desiccated before freezing).

Assessment of cultured media for desert moss crust by physiological responses.

The physiological responses of desert moss crusts under four artificial media (Beneck, Part, BG11, and Hogland) were investigated to evaluate the function of culture media during different culture periods. The results showed that the value of malondialdehyde (MDA) was at a maximum at 11d, on the contrary, chlorophyll-a, soluble protein, and soluble sugar were at a minimum. As the time increased, the value of MDA and soluble protein decreased faster in the Hogland, while the value of chlorophyll-a and soluble sugar increased. At the end of the culture period, the value of chlorophyll-a and soluble sugar was at a maximum in the Hogland, while the value of MDA and soluble protein was at a minimum. The results suggested that the Hogland medium had a promoting effect on the growth of desert moss crusts. The selected artificial cultivation medium towards wider and larger scale field applications of cultural desert biocrust was widely anticipated.

When facilitation meets clonal integration in forest canopies.

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.