The Moss Physcomitrium (Physcomitrella) patens: A Model Organism for Non-Seed Plants.

Since the discovery two decades ago that transgenes are efficiently integrated into the genome of Physcomitrella patens by homologous recombination, this moss has been a premier model system to study evolutionary developmental biology questions, stem cell reprogramming, and the biology of nonvascular plants. P patens was the first non-seed plant to have its genome sequenced. With this level of genomic information, together with increasing molecular genetic tools, a large number of reverse genetic studies have propelled the use of this model system. A number of technological advances have recently opened the door to forward genetics as well as extremely efficient and precise genome editing in P patens Additionally, careful phylogenetic studies with increased resolution have suggested that P patens emerged from within Physcomitrium Thus, rather than Physcomitrella patens, the species should be named Physcomitrium patens Here we review these advances and describe the areas where P patens has had the most impact on plant biology.

Morphological characterization of Hypnaceae (Bryopsida, Hypnales): Investigating four genera from Western Himalayas by using LM and SEM techniques.

The Hypnaceae is one of the largest and diversified family among the pleurocarpous mosses which consists of nearly 60 genera and 1,000 species around the world. In Pakistan, it is represented by 15 species and 8 genera. The current research article provides the detail morphological investigation of four different species (genera) of the family Hypnaceae that is, Gollania clarescens, Hypnum revolutum, Homomallium andoi, and Taxiphyllum taxirameum in the Western Himalayas, Pakistan. The research article reports the first SEM study of the leaf surfaces of the studied taxa, along with comprehensive morphological characters of the four species. The main objective of the research project is to present the comparative light and scanning electron microscopic study to discuss the morphology in detail because previously the family is just reported in different bryophyte checklists of the Western Himalayas. Based on results, morphological characters, micromorphological observations, qualitative and quantitative attributes, taxonomic keys for the studied taxa, distribution, and habitat are described. Stereoscope was used to analyze different qualitative characters, and light microscope was used to observe and measure the laminal and alar cells of the leaf. The SEM study reveals many important surface features like cell shape and cell wall. The laminal cells were mostly linear and elongated with thin cell walls. The specimens used for the SEM were air dried, so the laminal cells were somewhat constricted and concaved. The current study project will help to make the contribution in the taxonomy and morphology of this family.

Do mosses really exhibit so large distribution ranges? Insights from the integrative taxonomic study of the Lewinskya affinis complex (Orthotrichaceae, Bryopsida).

The strikingly lower number of bryophyte species, and in particular of endemic species, and their larger distribution ranges in comparison with angiosperms, have traditionally been interpreted in terms of their low diversification rates associated with a high long-distance dispersal capacity. This hypothesis is tested here with Lewinskya affinis (≡ Orthotrichum affine), a moss species widely spread across Europe, North and East Africa, southwestern Asia, and western North America. We tested competing taxonomic hypotheses derived from separate and combined analyses of multilocus sequence data, morphological characters, and geographical distributions. The best hypothesis, selected by a Bayes factor molecular delimitation analysis, established that L. affinis is a complex of no less than seven distinct species, including L. affinis s.str., L. fastigiata and L. leptocarpa, which were previously reduced into synonymy with L. affinis, and four new species. Discriminant analyses indicated that each of the seven species within L. affinis s.l. can be morphologically identified with a minimal error rate. None of these species exhibit a trans-oceanic range, suggesting that the broad distributions typically exhibited by moss species largely result from a taxonomic artefact. The presence of three sibling western North American species on the one hand, and four Old World sibling species on the other, suggests that there is a tendency for within-continent diversification rather than recurrent dispersal following speciation. The faster rate of diversification as compared to intercontinental migration reported here is in sharp contrast with earlier views of bryophyte species with wide ranges and low speciation rates.

Desiccation tolerance in the Antarctic moss Sanionia uncinata.

BACKGROUND: One of the most extreme environments on our planet is the Maritime Antarctic territory, due to its low-water availability, which restricts the development of plants. Sanionia uncinata Hedw. (Amblystegiaceae), the main colonizer of the Maritime Antarctic, has effective mechanisms to tolerate this environment. It has been described that the tolerance to desiccation is mediated by the hormone abscisic acid (ABA), antioxidants systems, accumulation of compatible solutes and proteins of the late embryogenesis abundant (LEA). However, to date, these mechanisms have not been described in S. uncinata. Therefore, in this work, we postulate that the tolerance to desiccation in the Antarctic moss S. uncinata is mediated by the accumulation of ABA, the osmolytes proline and glycine betaine, and dehydrins (an LEA class 11 proteins). To demonstrate our hypothesis, S. uncinata was subjected to desiccation for 24 h (loss in 95% of water content), and the effects on its physiological, photosynthetic, antioxidant and biochemical parameters were determined. RESULTS: Our results showed an accumulation of ABA in response to water loss, and the activation of protective responses that involves an increment in levels of proline and glycine betaine, an increment in the activity of antioxidant enzymes such as SOD, CAT, APX and POD, and the accumulation of dehydrins proteins. CONCLUSION: The results showed, suggest that S. uncinata is a  desiccation-tolerant moss, property mediated by high cellular plasticity regulated by ABA.

The long journey of Orthotrichum shevockii (Orthotrichaceae, Bryopsida): From California to Macaronesia.

Biogeography, systematics and taxonomy are complementary scientific disciplines. To understand a species' origin, migration routes, distribution and evolutionary history, it is first necessary to establish its taxonomic boundaries. Here, we use an integrative approach that takes advantage of complementary disciplines to resolve an intriguing scientific question. Populations of an unknown moss found in the Canary Islands (Tenerife Island) resembled two different Californian endemic species: Orthotrichum shevockii and O. kellmanii. To determine whether this moss belongs to either of these species and, if so, to explain its presence on this distant oceanic island, we combined the evaluation of morphological qualitative characters, statistical morphometric analyses of quantitative traits, and molecular phylogenetic inferences. Our results suggest that the two Californian mosses are conspecific, and that the Canarian populations belong to this putative species, with only one taxon thus involved. Orthotrichum shevockii (the priority name) is therefore recognized as a morphologically variable species that exhibits a transcontinental disjunction between western North America and the Canary Islands. Within its distribution range, the area of occupancy is limited, a notable feature among bryophytes at the intraspecific level. To explain this disjunction, divergence time and ancestral area estimation analyses are carried out and further support the hypothesis of a long-distance dispersal event from California to Tenerife Island.

Desiccation tolerance in the Antarctic moss Sanionia uncinata

BACKGROUND: One of the most extreme environments on our planet is the Maritime Antarctic territory, due to its low-water availability, which restricts the development of plants. Sanionia uncinata Hedw. (Amblystegiaceae), the main colonizer of the Maritime Antarctic, has effective mechanisms to tolerate this environment. It has been described that the tolerance to desiccation is mediated by the hormone abscisic acid (ABA), antioxidants systems, accumulation of compatible solutes and proteins of the late embryogenesis abundant (LEA). However, to date, these mechanisms have not been described in S. uncinata. Therefore, in this work, we postulate that the tolerance to desiccation in the Antarctic moss S. uncinata is mediated by the accumulation of ABA, the osmolytes proline and glycine betaine, and dehydrins (an LEA class 11 proteins). To demonstrate our hypothesis, S. uncinata was subjected to desiccation for 24 h (loss in 95% of water content), and the effects on its physiological, photosynthetic, antioxidant and biochemical parameters were determined. RESULTS: Our results showed an accumulation of ABA in response to water loss, and the activation of protective responses that involves an increment in levels of proline and glycine betaine, an increment in the activity of antioxidant enzymes such as SOD, CAT, APX and POD, and the accumulation of dehydrins proteins. CONCLUSION: The results showed, suggest that S. uncinata is a desiccation-tolerant moss, property mediated by high cellular plasticity regulated by ABA.

Molecular phylogeny of the genus Fissidens (Fissidentaceae, Bryophyta) and a refinement of the infrageneric classification.

The genus Fissidens (ca. 440 spp.) is one of the phylogenetically poorly studied groups of mosses (Bryophyta). While various classifications of this genus have been proposed, no attempt at a classification of the genus based on combined molecular and morphological evidence has been made. Here, we present for the first time a comprehensive phylogenetic tree consisting of 50 representatives of Fissidens, reconstructed using sequence data from chloroplast rbcL and rps4 genes. Ancestral state reconstructions provide three clear apomorphies within Fissidens: peristome teeth, limbidium and chromosome number. Based on the phylogeny and morphological reassessment, we recognize three subgenera, Pachyfissidens, Neoamblyothallia, and Fissidens. Subgenus Neoamblyothallia consists of two sections: Neoamblyothallia and Crispidium. Subgenus Fissidens consists of five sections: Fissidens, Polypodiopsis, Aloma, Areofissidens, and Semilimbidium. High diversity of the most derived sect. Semilimbidium in the tropics suggests that the evolutionary history of the genus is through adaptation and diversification in tropical regions.

Gene refashioning through innovative shifting of reading frames in mosses.

Early-diverging land plants such as mosses are known for their outstanding abilities to grow in various terrestrial habitats, incorporating tremendous structural and physiological innovations, as well as many lineage-specific genes. How these genes and functional innovations evolved remains unclear. In this study, we show that a dual-coding gene YAN/AltYAN in the moss Physcomitrella patens evolved from a pre-existing hemerythrin gene. Experimental evidence indicates that YAN/AltYAN is involved in fatty acid and lipid metabolism, as well as oil body and wax formation. Strikingly, both the recently evolved dual-coding YAN/AltYAN and the pre-existing hemerythrin gene might have similar physiological effects on oil body biogenesis and dehydration resistance. These findings bear important implications in understanding the mechanisms of gene origination and the strategies of plants to fine-tune their adaptation to various habitats.

The Complete Plastome Sequence of an Antarctic Bryophyte Sanionia uncinata (Hedw.) Loeske.

Organellar genomes of bryophytes are poorly represented with chloroplast genomes of only four mosses, four liverworts and two hornworts having been sequenced and annotated. Moreover, while Antarctic vegetation is dominated by the bryophytes, there are few reports on the plastid genomes for the Antarctic bryophytes. Sanionia uncinata (Hedw.) Loeske is one of the most dominant moss species in the maritime Antarctic. It has been researched as an important marker for ecological studies and as an extremophile plant for studies on stress tolerance. Here, we report the complete plastome sequence of S. uncinata, which can be exploited in comparative studies to identify the lineage-specific divergence across different species. The complete plastome of S. uncinata is 124,374 bp in length with a typical quadripartite structure of 114 unique genes including 82 unique protein-coding genes, 37 tRNA genes and four rRNA genes. However, two genes encoding the α subunit of RNA polymerase (rpoA) and encoding the cytochrome b6/f complex subunit VIII (petN) were absent. We could identify nuclear genes homologous to those genes, which suggests that rpoA and petN might have been relocated from the chloroplast genome to the nuclear genome.

Sequence analysis of the Hsp70 family in moss and evaluation of their functions in abiotic stress responses.

The 70-kD heat shock proteins (Hsp70s) are highly conserved molecular chaperones that play essential roles in cellular processes including abiotic stress responses. Physcomitrella patens serves as a representative of the first terrestrial plants and can recover from serious dehydration. To assess the possible relationship between P. patens Hsp70s and dehydration tolerance, we analyzed the P. patens genome and found at least 21 genes encoding Hsp70s. Gene structure and motif composition were relatively conserved in each subfamily. The intron-exon structure of PpcpHsp70-2 was different from that of other PpcpHsp70s; this gene exhibits several forms of intron retention, indicating that introns may play important roles in regulating gene expression. We observed expansion of Hsp70s in P. patens, which may reflect adaptations related to development and dehydration tolerance, and results mainly from tandem and segmental duplications. Expression profiles of rice, Arabidopsis and P. patens Hsp70 genes revealed that more than half of the Hsp70 genes were responsive to ABA, salt and drought. The presence of overrepresented cis-elements (DOFCOREZM and GCCCORE) among stress-responsive Hsp70s suggests that they share a common regulatory pathway. Moss plants overexpressing PpcpHsp70-2 showed salt and dehydration tolerance, further supporting a role in adaptation to land. This work highlights directions for future functional analyses of Hsp70s.

A phylotranscriptomic analysis of gene family expansion and evolution in the largest order of pleurocarpous mosses (Hypnales, Bryophyta).

The pleurocarpous mosses (i.e., Hypnanae) are a species-rich group of land plants comprising about 6,000 species that share the development of female sex organs on short lateral branches, a derived trait within mosses. Many of the families within Hypnales, the largest order of pleurocarpous mosses, trace their origin to a rapid radiation less than 100 million years ago, just after the rise of the angiosperms. As a result, the phylogenetic resolution among families of Hypnales, necessary to test evolutionary hypotheses, has proven difficult using one or few loci. We present the first phylogenetic inference from high-throughput sequence data (transcriptome sequences) for pleurocarpous mosses. To test hypotheses of gene family evolution, we built a species tree of 21 pleurocarpous and six acrocarpous mosses using over one million sites from 659 orthologous genes. We used the species tree to investigate the genomic consequences of the shift to pleurocarpy and to identify whether patterns common to other plant radiations (gene family expansion, whole genome duplication, or changes in the molecular signatures of selection) could be observed. We found that roughly six percent of all gene families have expanded in the pleurocarpous mosses, relative to acrocarpous mosses. These gene families are enriched for several gene ontology (GO) terms, including interaction with other organisms. The increase in copy number coincident with the radiation of Hypnales suggests that a process such as whole genome duplication or a burst of small-scale duplications occurred during the diversification. In over 500 gene families we found evidence of a reduction in purifying selection. These gene families are enriched for several terms in the GO hierarchy related to "tRNA metabolic process." Our results reveal candidate genes and pathways that may be associated with the transition to pleurocarpy, illustrating the utility of phylotranscriptomics for the study of molecular evolution in non-model species.

A Transcriptome Atlas of Physcomitrella patens Provides Insights into the Evolution and Development of Land Plants.

Identifying the genetic mechanisms that underpin the evolution of new organ and tissue systems is an aim of evolutionary developmental biology. Comparative functional genetic studies between angiosperms and bryophytes can define those genetic changes that were responsible for developmental innovations. Here, we report the generation of a transcriptome atlas covering most phases in the life cycle of the model bryophyte Physcomitrella patens, including detailed sporophyte developmental progression. We identified a comprehensive set of sporophyte-specific transcription factors, and found that many of these genes have homologs in angiosperms that function in developmental processes such as flowering and shoot branching. Deletion of the PpTCP5 transcription factor results in development of supernumerary sporangia attached to a single seta, suggesting that it negatively regulates branching in the moss sporophyte. Given that TCP genes repress branching in angiosperms, we suggest that this activity is ancient. Finally, comparison of P. patens and Arabidopsis thaliana transcriptomes led us to the identification of a conserved core of transcription factors expressed in tip-growing cells. We identified modifications in the expression patterns of these genes that could account for developmental differences between P. patens tip-growing cells and A. thaliana pollen tubes and root hairs.

Population structure and genetic diversity in the nannandrous moss Homalothecium lutescens: does the dwarf male system facilitate gene flow?

BACKGROUND: Nannandry is a sexual system where males ("dwarf males") are much smaller than the conspecific females. Dwarf males occur in a wide range of unrelated organisms but the evolutionary advantages of this condition are poorly understood. The dwarf male sexual system results in differences in the mode of dispersal and establishment as well as the life span between males and females. Such differences must have profound effects on the population dynamics and genetic structures. We have studied four populations of the nannandrous moss Homalothecium lutescens in southern Sweden. We genotyped dwarf males and female shoots with the aim of describing the genetic diversity and structure of the populations. RESULTS: Dwarf males were most related to their host shoot, then their colony (within 0.5 m(2)) and then the rest of the population, which suggests restricted spore dispersal. However, a few dwarf males in each population appeared to originate from other colonies and sometimes even other populations. Genetic diversity of dwarf males was generally high but showed no tendency to be consistently higher or lower than female genetic diversity within the four populations. CONCLUSIONS: Although most dwarf males have local origin, sporadic dispersal events occur. The ability of the dwarf males to establish in high numbers in mature colonies facilitates gene flow between populations as well as increases the potential to accumulate genetic diversity within populations.

[BIOINFORMATICS COMPARISON OF HUMAN AND PLANT PHOSPHATOMES].

The study presents the results of bioinformatic comparison of protein phosphatases from higher plants and human phosphatom (150 sequences). Based on sequence and profile comparison with known catalytic domains, 204 plant homologues from Physcomitrella patens and Arabidopsis thaliana where selected. Clustering of joint group of plant and mammalian protein phosphatases revealed fundamental differences in plant and human phosphatomes. At the same time, it was shown significant differences in the set of protein phosphatases in P. patens, A. thaliana, and such monocots as Orysa saliva and Zea mays.

Impacts of Environmental Heterogeneity on Moss Diversity and Distribution of Didymodon (Pottiaceae) in Tibet, China.

Tibet makes up the majority of the Qinghai-Tibet Plateau, often referred to as the roof of the world. Its complex landforms, physiognomy, and climate create a special heterogeneous environment for mosses. Each moss species inhabits its own habitat and ecological niche. This, in combination with its sensitivity to environmental change, makes moss species distribution a useful indicator of vegetation alteration and climate change. This study aimed to characterize the diversity and distribution of Didymodon (Pottiaceae) in Tibet, and model the potential distribution of its species. A total of 221 sample plots, each with a size of 10 × 10 m and located at different altitudes, were investigated across all vegetation types. Of these, the 181 plots in which Didymodon species were found were used to conduct analyses and modeling. Three noteworthy results were obtained. First, a total of 22 species of Didymodon were identified. Among these, Didymodon rigidulus var. subulatus had not previously been recorded in China, and Didymodon constrictus var. constrictus was the dominant species. Second, analysis of the relationships between species distributions and environmental factors using canonical correspondence analysis revealed that vegetation cover and altitude were the main factors affecting the distribution of Didymodon in Tibet. Third, based on the environmental factors of bioclimate, topography and vegetation, the distribution of Didymodon was predicted throughout Tibet at a spatial resolution of 1 km, using the presence-only MaxEnt model. Climatic variables were the key factors in the model. We conclude that the environment plays a significant role in moss diversity and distribution. Based on our research findings, we recommend that future studies should focus on the impacts of climate change on the distribution and conservation of Didymodon.

Appearance and elaboration of the ethylene receptor family during land plant evolution.

Ethylene is perceived following binding to endoplasmic reticulum-localized receptors, which in Arabidopsis thaliana, include ETR1, ERS1, EIN4, ETR2, and ERS2. These receptors fall into two subfamilies based on conservation of features within their histidine kinase domain. Subfamily 1 contains ETR1 and ERS1 whereas subfamily 2 contains EIN4, ETR2, and ERS2. Because ethylene receptors are found only in plants, this raises questions of when each receptor evolved. Here it is shown that subfamily 1 receptors encoded by a multigene family are present in all charophytes examined, these being most homologous to ETR1 based on their evolutionary relationship as well as containing histidine kinase and receiver domains. In charophytes and Physcomitrella patens, one or more gene family members contain the intron characteristic of subfamily 2 genes, indicating the first step in subfamily 2 receptor evolution. ERS1 homologs appear in basal angiosperm species after Amborella trichopoda and, in some early and basal angiosperm species and monocots in general, it is the only subfamily 1 receptor present. Distinct EIN4 and ETR2 homologs appear only in core eudicots and ERS2 homologs appear only in the Brassicaceae, suggesting it is the most recent receptor to evolve. These findings show that a subfamily 1 receptor had evolved and a subfamily 2 receptor had begun to evolve in plants prior to the colonization of land and only these two existed up to the appearance of the first basal angiosperm. The appearance of ERS2 in the Brassicaceae suggests ongoing evolution of the ethylene receptor family.

Transcriptomic evidence for the evolution of shoot meristem function in sporophyte-dominant land plants through concerted selection of ancestral gametophytic and sporophytic genetic programs.

Alternation of generations, in which the haploid and diploid stages of the life cycle are each represented by multicellular forms that differ in their morphology, is a defining feature of the land plants (embryophytes). Anciently derived lineages of embryophytes grow predominately in the haploid gametophytic generation from apical cells that give rise to the photosynthetic body of the plant. More recently evolved plant lineages have multicellular shoot apical meristems (SAMs), and photosynthetic shoot development is restricted to the sporophyte generation. The molecular genetic basis for this evolutionary shift from gametophyte-dominant to sporophyte-dominant life cycles remains a major question in the study of land plant evolution. We used laser microdissection and next generation RNA sequencing to address whether angiosperm meristem patterning genes expressed in the sporophytic SAM of Zea mays are expressed in the gametophytic apical cells, or in the determinate sporophytes, of the model bryophytes Marchantia polymorpha and Physcomitrella patens. A wealth of upregulated genes involved in stem cell maintenance and organogenesis are identified in the maize SAM and in both the gametophytic apical cell and sporophyte of moss, but not in Marchantia. Significantly, meiosis-specific genetic programs are expressed in bryophyte sporophytes, long before the onset of sporogenesis. Our data suggest that this upregulated accumulation of meiotic gene transcripts suppresses indeterminate cell fate in the Physcomitrella sporophyte, and overrides the observed accumulation of meristem patterning genes. A model for the evolution of indeterminate growth in the sporophytic generation through the concerted selection of ancestral meristem gene programs from gametophyte-dominant lineages is proposed.

Molecular phylogenetic study and expression analysis of ATP-binding cassette transporter gene family in Oryza sativa in response to salt stress.

ATP-binding cassette (ABC) transporter is a large gene superfamily that utilizes the energy released from ATP hydrolysis for transporting myriad of substrates across the biological membranes. Although many investigations have been done on the structural and functional analysis of the ABC transporters in Oryza sativa, much less is known about molecular phylogenetic and global expression pattern of the complete ABC family in rice. In this study, we have carried out a comprehensive phylogenetic analysis constructing neighbor-joining and maximum-likelihood trees based on various statistical methods of different ABC protein subfamily of five plant lineages including Chlamydomonas reinhardtii (green algae), Physcomitrella patens (moss), Selaginella moellendorffii (lycophyte), Arabidopsis thaliana (dicot) and O. sativa (monocot) to explore the origin and evolutionary patterns of these ABC genes. We have identified several conserved motifs in nucleotide binding domain (NBD) of ABC proteins among all plant lineages during evolution. Amongst the different ABC protein subfamilies, 'ABCE' has not yet been identified in lower plant genomes (algae, moss and lycophytes). The result indicated that gene duplication and diversification process acted upon these genes as a major operative force creating new groups and subgroups and functional divergence during evolution. We have demonstrated that rice ABCI subfamily consists of only half size transporters that represented highly dynamic members showing maximum sequence variations among the other rice ABC subfamilies. The evolutionary and the expression analysis contribute to a deep insight into the evolution and diversity of rice ABC proteins and their roles in response to salt stress that facilitate our further understanding on rice ABC transporters.

Molecular evidence for convergent evolution and allopolyploid speciation within the Physcomitrium-Physcomitrella species complex.

BACKGROUND: The moss Physcomitrella patens (Hedw.) Bruch & Schimp. is an important experimental model system for evolutionary-developmental studies. In order to shed light on the evolutionary history of Physcomitrella and related species within the Funariaceae, we analyzed the natural genetic diversity of the Physcomitrium-Physcomitrella species complex. RESULTS: Molecular analysis of the nuclear single copy gene BRK1 reveals that three Physcomitrium species feature larger genome sizes than Physcomitrella patens and encode two expressed BRK1 homeologs (polyploidization-derived paralogs), indicating that they may be allopolyploid hybrids. Phylogenetic analyses of BRK1 as well as microsatellite simple sequence repeat (SSR) data confirm a polyphyletic origin for three Physcomitrella lineages. Differences in the conservation of mitochondrial editing sites further support hybridization and cryptic speciation within the Physcomitrium-Physcomitrella species complex. CONCLUSIONS: We propose a revised classification of the previously described four subspecies of Physcomitrella patens into three distinct species, namely Physcomitrella patens, Physcomitrella readeri and Physcomitrella magdalenae. We argue that secondary reduction of sporophyte complexity in these species is due to the establishment of an ecological niche, namely spores resting in mud and possible spore dispersal by migratory birds. Besides the Physcomitrium-Physcomitrella species complex, the Funariaceae are host to their type species, Funaria hygrometrica, featuring a sporophyte morphology which is more complex. Their considerable developmental variation among closely related lineages and remarkable trait evolution render the Funariaceae an interesting group for evolutionary and genetic research.

Genotypic variation in a foundation tree (Populus tremula L.) explains community structure of associated epiphytes.

Community genetics hypothesizes that within a foundation species, the genotype of an individual significantly influences the assemblage of dependent organisms. To assess whether these intra-specific genetic effects are ecologically important, it is required to compare their impact on dependent organisms with that attributable to environmental variation experienced over relevant spatial scales. We assessed bark epiphytes on 27 aspen (Populus tremula L.) genotypes grown in a randomized experimental array at two contrasting sites spanning the environmental conditions from which the aspen genotypes were collected. We found that variation in aspen genotype significantly influenced bark epiphyte community composition, and to the same degree as environmental variation between the test sites. We conclude that maintaining genotypic diversity of foundation species may be crucial for conservation of associated biodiversity.