Evolution of sexual systems, dispersal strategies and habitat selection in the liverwort genus Radula

? Shifts in sexual systems are among the most common and important transitions in plants and are correlated with a suite of life-history traits. The evolution of sexual systems and their relationships to gametophyte size, sexual and asexual reproduction, and epiphytism are examined here in the liverwort genus Radula. ? The sequence of trait acquisition and the phylogenetic correlations between those traits was investigated using comparative methods. ? Shifts in sexual systems recurrently occurred from dioecy to monoecy within facultative epiphyte lineages. Production of specialized asexual gemmae was correlated to neither dioecy nor strict epiphytism. ? The significant correlations among life-history traits related to sexual systems and habitat conditions suggest the existence of evolutionary trade-offs. Obligate epiphytes do not produce gemmae more frequently than facultative epiphytes and disperse by whole gametophyte fragments, presumably to avoid the sensitive protonemal stage in a habitat prone to rapid changes in moisture availability. As dispersal ranges correlate with diaspore size, this reinforces the notion that epiphytes experience strong dispersal limitations. Our results thus provide the evolutionary complement to metapopulation, metacommunity and experimental studies demonstrating trade-offs between dispersal distance, establishment ability, and life-history strategy, which may be central to the evolution of reproductive strategies in bryophytes.     

Reproductive strategy, clonal structure and genetic diversity in populations of the aquatic macrophyte Sparganium emersum in river systems

Many aquatic and riparian plant species are characterized by the ability to reproduce both sexually and asexually. Yet, little is known about how spatial variation in sexual and asexual reproduction affects the genotypic diversity within populations of aquatic and riparian plants. We used six polymorphic microsatellites to examine the genetic diversity within and differentiation among 17 populations (606 individuals) of Sparganium emersum, in two Dutch-German rivers. Our study revealed a striking difference between rivers in the mode of reproduction (sexual vs. asexual) within S. emersum populations. The mode of reproduction was strongly related to locally reigning hydrodynamic conditions. Sexually reproducing populations exhibited a greater number of multilocus genotypes compared to asexual populations. The regional population structure suggested higher levels of gene flow among sexually reproducing populations compared to clonal populations. Gene flow was mainly mediated via hydrochoric dispersal of generative propagules (seeds), impeding genetic differentiation among populations even over river distances up to 50 km. Although evidence for hydrochoric dispersal of vegetative propagules (clonal plant fragments) was found, this mechanism appeared to be relatively less important. Bayesian-based assignment procedures revealed a number of immigrants, originating from outside our study area, suggesting intercatchment plant dispersal, possibly the result of waterfowl-mediated seed dispersal. This study demonstrates how variation in local environmental conditions in river systems, resulting in shifting balances of sexual vs. asexual reproduction within populations, will affect the genotypic diversity within populations. This study furthermore cautions against generalizations about dispersal of riparian plant species in river systems.     

Interactions among species with contrasting dispersal modes explain distributions for epiphytic lichens

Understanding how the biodiversity response to climate change will be modified at ecological scales, e.g. by species interactions, is a major challenge. Lichen epiphytes – the close interdependent relationship between a heterotrophic fungus and photosynthetic partner (photobiont) – are used here to explore how interaction regimes (between lichen species, and between lichens and their photobionts) explain distribution patterns along spatial climatic gradients. To do this we tested field evidence for the ‘core‐fringe hypothesis’, which proposes a facilitative interaction; sexually‐reproducing and spore‐dispersed lichens with a requirement for resynthesis with a compatible photobiont (Nostoc) are facilitated by the prior establishment of asexual lichens which disperse both the fungus and photobiont together. We used two closely related Nephroma species which differ in their reproductive mode – N. laevigatum (sexual spore‐dispersed) and N. parile (asexual) – and compared their occurrence along a bioclimatic gradient to local habitat factors, including the co‐occurrence of asexual lichens which have shared specificity for compatible Nostoc genotypes. The results showed that: 1) N. laevigatum is significantly more likely to occur on trees that have already been colonised by asexual lichens with shared specificity for Nostoc, supporting the core‐fringe hypothesis, while 2) N. parile is independent of this association (strengthening the core‐fringe hypothesis), with its response to a precipitation gradient modified by microhabitat factors. This positive test for the core‐fringe hypothesis demonstrates how interaction regimes can fundamentally alter expectations under climate change. There is an assumption that spore‐dispersed lichen species could more easily track their suitable bioclimatic space through fragmented habitat, compared to asexual species with larger and heavier propagules. However, the establishment of spore‐dispersed lichen epiphytes into new habitat may be limited by the dispersal rates of asexual species, which act as key facilitators.     

The reproductive biology of Polytrichum formosum: clonal structure and paternity revealed by microsatellites

Using highly polymorphic microsatellite markers, we assessed clonal structure and paternity in a population of the bryophyte species Polytrichum formosum. Identical multilocus genotypes of individual shoots were almost never observed in spatially separated cushions, but were found to be highly clustered within moss cushions. Therefore, asexual reproduction through dispersal of gametophyte fragments is not very important in P. formosum. However, asexual reproduction on a very localized scale through vegetative growth of genets (branching of gametophytes via clonal growth of rhizomes) is very extensive. The patchy spatial distribution of genets and the absence of intermingling among genets suggest that this species follows a 'phalanx' clonal growth strategy. Vegetative proliferation of genets will increase their size, and, consequently, will have considerable fitness consequences for individuals in terms of increased genet longevity and reproductive output. Although paternity analysis of sporophytes confirmed male genet size, i.e. gamete production, to be an important determinant of male reproductive fitness, it also showed that the spatial distance to female genets is the predominant factor that governs male reproductive success. Moreover, we showed that male gamete dispersal distances in P. formosum are much further than generally assumed, and are in the order of metres rather than centimetres. Combining the findings, we conclude that the high genotypic diversity observed for this facultatively clonal species is most likely explained by a preponderance of sexual reproduction over clonal reproduction.     

The relative importance of sexual and asexual reproduction in the spread of Spartina alterniflora using a spatially explicit individual-based model

This study investigates a spatially explicit, individual-based model for simulating the spread of invasive smooth cordgrass (Spartina alterniflora) in Yancheng coastal wetlands from 1995 to 2010. The model, which considers the landscape heterogeneity and changes detected by remote sensing, also reveals the relative importance of sexual and asexual reproduction in the spread by global sensitivity analysis. The model was verified as suitable for simulating the range expansion of S. alterniflora. The results show that: (1) although seedling recruitment is low, it significantly contributes to the range expansion of S. alterniflora. Removing sexual propagation greatly reduces the expansion rate. Rapid expansion requires both sexual and asexual reproduction; (2) in the global sensitivity analysis, the most significant affecters of S. alterniflora invasion were seed dispersal distance, adult survival rate and asexual recruitment survival rate. Sexual propagation contributes much more significantly to quick range expansion than asexual reproduction, but asexual reproduction is the main source of recruitment. Invasion control strategies should target a single reproduction mode. Here, limiting the germination and dispersal of seeds is suggested as a realistic strategy for controlling and managing invasion by this species.     

An experimental demonstration of the cost of sex and a potential resource limitation on reproduction in the moss Pterygoneurum (Pottiaceae)

The cost of sexual reproduction is incurred when the current reproductive episode contributes to a a decline in future plant performance. To test the hypotheses that a trade-off exists between current sexual reproduction and subsequent clonal regeneration and that resources limit reproduction and regeneration, plants of the widespread moss Pterygoneurum ovatum were subjected to induced sporophytic abortion, upper leaf removal, and nutrient amendment treatments. Sexually reproducing plants were slower or less likely to produce regenerative structures (protonemata or shoots) and produced fewer regenerative tissue areas or structures. The ability and the timeline to reproduce sexually and regenerate clonally were unaffected by an inorganic nutrient amendment. However, when leaves subtending the sporophyte were removed, the sporophytes were less likely to mature, tended to take a longer time to mature, and were smaller compared to sporophytes from shoots with a full complement of upper leaves. Our findings indicate that plants investing in sexual reproduction suffer a cost of decreased clonal regeneration and indicate that sporophyte maturation is resource-limited, with upper leaves contributing to the nutrition of the sporophyte. This study represents only the second explicit experimental demonstration of a trade-off between sexual and asexual reproduction in bryophytes.     

Notes on African Mosses. I. Andreaea camerunensis sp.nov. and other Mosses mostly from Nigeria and the British Cameroons

Abstract

Reproductive ecological traits such as success of fertilization, partitioning of sexes, the relative success of sexual versus asexual reproduction and dispersal distances are likely to considerably influence genetic structure within and among plant populations. In the liverwort Mannia fragrans both sexual and asexual reproduction can be frequently observed: sporophytes are produced abundantly every year and asexual propagation by fragmentation of thalli is also common. The aim of this study was to use ecological and molecular methods (ISSR markers) to separate the role of sexual and asexual components in shaping the partitioning of genetic variability within and among populations. In addition to genetic analyses conducted seasonally, sex expression and fertilization rates, sex ratios, regeneration from vegetative fragments and outcrossing was estimated in 3 populations of the species. Sex expression rates were high and, in spite of strongly female biased sex ratios, high fertilization rates were detected. However, capacity for regeneration from fragments was also high. Despite frequent spore production genetic diversity was low within populations. This is probably the result of the predominance of asexual reproduction s. 1. including crossing between genetically identical plants. Although recombination and mutation occasionally generates new haplotypes, these have little chance to spread because of the large spores mainly falling into their own patch, where chances for germination are low. Due to small size and isolation of the populations, genetic drift is likely to eliminate these haplotypes. Remote populations differed significantly, each being dominated by a few clones, reflecting negligible gene flow among them. Differences among individual populations can partly be related to differences in their reproductive behaviour and degree of isolation.     

The eco-evolutionary importance of reproductive system variation in the macroalgae: Freshwater reds as a case study

The relative frequency of sexual versus asexual reproduction governs the distribution of genetic diversity within and among populations. Most studies on the consequences of reproductive variation focus on the mating system (i.e., selfing vs. outcrossing) of diploid-dominant taxa (e.g., angiosperms), often ignoring asexual reproduction. Although reproductive systems are hypothesized to be correlated with life-cycle types, variation in the relative rates of sexual and asexual reproduction remains poorly characterized across eukaryotes. This is particularly true among the three major lineages of macroalgae (green, brown, and red). The Rhodophyta are particularly interesting, as many taxa have complex haploid–diploid life cycles that influence genetic structure. Though most marine reds have separate sexes, we show that freshwater red macroalgae exhibit patterns of switching between monoicy and dioicy in sister taxa that rival those recently shown in brown macroalgae and in angiosperms. We advocate for the investigation of reproductive system evolution using freshwater reds, as this will expand the life-cycle types for which these data exist, enabling comparative analyses broadly across eukaryotes. Unlike their marine cousins, species in the Batrachospermales have macroscopic gametophytes attached to filamentous, often microscopic sporophytes. While asexual reproduction through monospores may occur in all freshwater reds, the Compsopogonales are thought to be exclusively asexual. Understanding the evolutionary consequences of selfing and asexual reproduction will aid in our understanding of the evolutionary ecology of all algae and of eukaryotic evolution generally.     

How to go extinct by mating too much: population consequences of male mate choice and efficiency in a sexual–asexual species complex

Selection acting on individuals is not predicted to maximize population persistence, yet examples that explicitly quantify conflicts between individual and population level benefits are scarce. One such conflict occurs over sexual reproduction because of the cost of sex: sexual populations that suffer the cost of producing males have only half the growth rate compared to asexuals. Male behaviour can additionally impact population dynamics in a variety of ways, and here we study an example where the impact is unusually clear: the riddle of persistence of sperm‐dependent sexual–asexual species complexes. Here, a sexually reproducing host species coexists with an ameiotically reproducing all‐female sperm parasite. Sexual–asexual coexistence should not be stable because the proportion of asexually reproducing females will rapidly increase and the relative abundance of the sexually reproducing host species will decline. A severe shortage of males will lead to sperm limitation for sexual and asexual females and the system collapses. Male mate choice could reduce the reproductive potential of the asexual species and thus potentially prevent the collapse. In the gynogenetic (sperm‐dependent parthenogenetic) Amazon molly Poecilia formosa and its host (P. latipinna or P. mexicana), males discriminate against asexual females to some extent. Using a population‐dynamical model, we examine the population dynamics of this species complex with varying strengths of male discrimination ability and efficiency with which they locate females and produce sperm. The sexual species would benefit from stronger discrimination, thus preventing being displaced by the asexual females. However, males would be required to evolve preferences that are probably too strong to be purely based upon selection acting on individuals. We conclude that male behaviour does not fully prevent but delays extinction, yet this is highly relevant because low local extinction rates strongly promote coexistence as a metapopulation.     

中国藓类植物无性繁殖体的初步观察

无性生殖在苔藓植物的生活史中起着重要的作用,并且常通过各种无性繁殖体来完成。无性繁殖体的形态常被用来辅助鉴定一些不育的藓类植物。本文通过对38种藓类植物的无性繁殖体进行显微观察,结果显示：无性繁殖体在不同的藓类植物之间已经过多次演化;无性繁殖体的形态在种内是相对稳定的,且与其着生位置、配子体的分枝方式密切相关,而与植物的系统位置以及生境并无直接的关系;无性繁殖体的颜色与其胞壁的厚度以及表面纹饰密切相关。此外,无性繁殖体的产生常常与假根、原丝体有共存关系。在研究中发现,藓类植物的无性繁殖体主要包括原丝体芽胞、无性芽胞（叶生芽胞、中肋芽胞、枝生芽胞）、芽体、假根芽胞和假根状块茎;其中原丝体芽胞和无性芽胞最为常见。     

Population subdivision in Australian temperate marine invertebrates: Larval connections versus historical factors*

This paper reviews the use of genetic data, in combination with manipulative experimentation, to infer the mode of reproduction and the extent and directionality of dispersal for a range of Australian temperate marine invertebrates. Local populations of obligately sexually reproducing species have been inferred to be strongly interconnected by larval dispersal, over distances of thousands of kilometres. Their larvae may be subject to strong post-settlement selection, but this selection is independent of obvious geographic or intertidal gradients. Within local populations selection may therefore result in apparently chaotic genetic patchiness which is eliminated by the effects of sexual reproduction and the widespread dispersal and mixing of the colonizing larvae of each generation. In partial contrast, local populations of species which rely on asexual reproduction for the maintenance of populations show evidence of similar larval connections, but no recent settlement of their sexually generated larvae has been demonstrated. The apparent connectedness of these populations may reflect either historical events or a more episodic pattern of settlement by sexually generated larvae. Local populations of these species are more highly differentiated as a result of the continued asexual replication of a limited number of genotypes. In one of these species, reciprocal transplantation of the clones within and among populations has revealed that resident clones can be highly locally adapted (as reflected by much higher asexual fecundity), which implies that selection is an important determinant of the composition of local populations. Nevertheless, the failure to detect continuing sexual recruitment into these populations obscures the evolutionary significance of this finding.     

The evolutionary ecology (evo-eco) of plant asexual reproduction

The different modes of asexual plant reproduction are reviewed in the context of the evolutionary ecology (evo-eco) of retaining old (and colonizing new) habitats and exploring new ones. The modes of asexual reproduction are contrasted between the green algae and the nonvascular and vascular land plants (bryophytes and tracheophytes, respectively). The chronology of their appearance is also traced in the fossil record. Among bryophytes, the indeterminate growth of protonema and the subsequent fragmentation of gametophytes provide a successful asexual reproductive stratagem, one that is paralleled by indeterminate rhizomaty and subsequent fragmentation of tracheophyte sporophytes. Self-compatible (autogamous) bisexual (monoicous) gametophytes capable of producing viable homozygous sporophytes offer a method for colonizing new sites by means of a single spore, a stratagem that is paralleled in tracheophyte sporophytes by the co-option of sexual reproductive structures (e.g., flowers) to produce diploid embryos asexually (e.g., variants of apomixis such as nucellar embryony). A review of the fossil record and the biology of extant taxa show that some of these modes of asexual reproduction are ancient and present in the green ancestors of the land plants (i.e., the charophycean algae). An overview of the data indicates that asexual reproduction provides a bet-hedging stratagem that can obviate some of the many costs of sexual reproduction, and preserve well-adapted genomes within previously occupied sites.     

LOWER MITE INFESTATIONS IN AN ASEXUAL GECKO COMPARED WITH ITS SEXUAL ANCESTORS

What advantage do sexually reproducing organisms gain from their mode of reproduction that compensates for their twofold loss in reproductive rate relative to their asexual counterparts? One version of the Red Queen hypothesis suggests that selective pressure from parasites is strongest on the most common genotype in a population, and thus genetically identical clonal lineages are more vulnerable to parasitism over time than genetically diverse sexual lineages. Our surveys of the ectoparasites of an asexual gecko and its two sexual ancestral species show that the sexuals have a higher prevalence, abundance, and mean intensity of mites than asexuals sharing the same habitat. Our experimental data indicate that in one sexual/asexual pair this pattern is at least partly attributable to higher attachment rates of mites to sexuals. Such a difference may occur as a result of exceptionally high susceptibility of the sexuals to mites because of their low genetic diversity (relative to other more-outbred sexual species) and their potentially high stress levels, or as a result of exceptionally low susceptibility of the asexuals to mites because of their high levels of heterozygosity.     

Effects of forest canopy on habitat selection in treefrogs and aquatic insects: implications for communities and metacommunities

The specific dispersal/colonization strategies used by species to locate and colonize habitat patches can strongly influence both community and metacommunity structure. Habitat selection theory predicts nonrandom dispersal to and colonization of habitat patches based on their quality. We tested whether habitat selection was capable of generating patterns of diversity and abundance across a transition of canopy coverage (open and closed canopy) and nutrient addition by investigating oviposition site choice in two treefrog species (Hyla) and an aquatic beetle (Tropisternus lateralis), and the colonization dynamics of a diverse assemblage of aquatic insects (primarily beetles). Canopy cover produced dramatic patterns of presence/absence, abundance, and species richness, as open canopy ponds received 99.5% of propagules and 94.6% of adult insect colonists. Nutrient addition affected only Tropisternus oviposition, as females oviposited more egg cases at higher nutrient levels, but only in open canopy ponds. The behavioral partitioning of aquatic landscapes into suitable and unsuitable habitats via habitat selection behavior fundamentally alters how communities within larger ecological landscapes (metacommunities) are linked by dispersal and colonization.     

Sexual reproduction prevails in a world of structured resources in short supply

We present a model for the maintenance of sexual reproduction based on the availability of resources, which is the strongest factor determining the growth of populations. The model compares completely asexual species to species that switch between asexual and sexual reproduction (sexual species). Key features of the model are that sexual reproduction sets in when resources become scarce, and that at a given place only a few genotypes can be present at the same time. We show that under a wide range of conditions the sexual species outcompete the asexual ones. The asexual species win only when survival conditions are harsh and death rates are high, or when resources are so little structured or consumer genotypes are so manifold that all resources are exploited to the same extent. These conditions largely represent the conditions in which sexuals predominate over asexuals in the field.     

Reproductive strategies in lichens

Lichens employ two divergent strategies of reproduction, a sexual method through spore propagation and a derivative asexual strategy based on systems perpetuating the symbiosis by the dissemination of specialized diaspores. The distribution of these pathways in 24 families in the order Lecanorales is presented. Fruticose and foliose life forms have a greater representation of vegetative diaspores than crustose forms. It is hypothesized that chemical evolution occurred prior to the development of secondary asexual reproductive mechanisms. The derived vegetative morphologies have apparently outlasted their sexual ancestors, although a number of closely related "species pairs" are known, indicating the joint survival of ancestral sexual lines with secondary asexual offspring. Species using a vegetative strategy often have a greater world distribution than their sexual counterparts in foliose and fruticose groups. No clear pattern is apparent in crustose groups. The advantages of vegetative strategies are a greater survival for propagules and rapid invasion of new habitats. The separation of these strategies and the evolution of soredia and isidia represents a gradient in the evolution of taxa.     

基于个体的空间显性模型和遥感技术模拟入侵植物扩张机制

基于个体的空间显性模型和遥感技术,以互花米草为例,模拟了自1997到2010年的种群扩张动态,揭示了土地利用变化与潮间带高程的影响;并通过全局敏感性分析揭示了种子扩散、成体存活率、有性和无性繁殖等种群统计学特征对互花米草种群扩张的相对重要性。研究结果发现:1)有性繁殖与无性繁殖共同决定互花米草种群快速扩张;2)潮间带高程和土地利用变化显著影响模型预测的精度,对互花米草种群扩张有非常重要的影响;3)成体存活率与种子长距离扩散是影响互花米草种群扩张速度最重要的因素;无性繁殖比有性繁殖对种群扩张的影响更大;种子长距离扩散比本地扩散更为重要,同时,小概率的种子长距离扩散事件对种群扩张有非常重要的影响。为了经济有效地控制外来入侵植物的扩张,应该抑制种子的长距离扩散和移除种子长距离扩散形成的位于入侵前沿的小斑块。     

Restriction of sexual reproduction in the moss Racomitrium lanuginosum along an elevational gradient

1. Terrestrial plant populations located at the margins of species’ distributions often display reduced sexual reproduction and an increased reliance on asexual reproduction. One hypothesis to explain this phenomenon is that the decline is associated with environmental effects on the energetic costs to produce reproductive organs.
2. In order to clarify the changing processes of sexual reproduction along an elevational gradient, we investigated the sexual reproductive parameters, such as the number of sporophytes and gametangia, in Racomitrium lanuginosum, a dioicous moss found on Mt. Fuji.
3. Matured sporophytes were present only below 3,000 m, and the number of sporophytes per shoot tended to be lower at higher elevation habitats. The numbers of male inflorescences per shoot and antheridia per inflorescence and shoot significantly decreased with increasing elevation. In contrast, the numbers of female inflorescences per shoot and archegonia per inflorescence and shoot varied little across elevations.
4. Synthesis. Our results suggest that the reasons for this limitation are assumed to be limitations in sporophyte development that result in abortion, and the spatial segregation between males and females. Possible reasons for the abortion of sporophytes are the inhibitory effects of low air temperature, a shortened growth period, and winter environmental conditions at higher elevations. Remarkable differences between male and female on various reproductive parameters found in this study are thought to affect the mode of sexual reproduction under the harsh environment. These differences between males and females may be caused by differences in the costs of production and development of gametangia, sensitivity to environmental stressors, and phenological patterns.

     

Metacommunity patterns in larval odonates

The growth of metacommunity ecology as a subdiscipline has increased interest in how processes at different spatial scales structure communities. However, there is still a significant knowledge gap with respect to relating the action of niche- and dispersal-assembly mechanisms to observed species distributions across gradients. Surveys of the larval dragonfly community (Odonata: Anisoptera) in 57 lakes and ponds in southeast Michigan were used to evaluate hypotheses about the processes regulating community structure in this system. We considered the roles of both niche- and dispersal-assembly processes in determining patterns of species richness and composition across a habitat gradient involving changes in the extent of habitat permanence, canopy cover, area, and top predator type. We compared observed richness patterns and species distributions in this system to patterns predicted by four general community models: species sorting related to adaptive trade-offs, a developmental constraints hypothesis, dispersal assembly, and a neutral community assemblage. Our results supported neither the developmental constraints nor the neutral-assemblage models. Observed patterns of richness and species distributions were consistent with patterns expected when adaptive tradeoffs and dispersal-assembly mechanisms affect community structure. Adaptive trade-offs appeared to be important in limiting the distributions of species which segregate across the habitat gradient. However, dispersal was important in shaping the distributions of species that utilize habitats with a broad range of hydroperiods and alternative top predator types. Our results also suggest that the relative importance of these mechanisms may change across this habitat gradient and that a metacommunity perspective which incorporates both niche- and dispersal-assembly processes is necessary to understand how communities are organized. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Species richness patterns and metapopulation processes – evidence from epiphyte communities in boreo-nemoral forests

For several epiphyte species, dispersal limitation and metapopulation dynamics have been suggested. We studied the relative importance of local environmental conditions and spatial aggregation of species richness of facultative and obligate epiphytic bryophytes and lichens within two old‐growth forests in eastern Sweden. The effect of the local environment was analyzed using generalized linear models (GLM). We tested whether species richness was spatially structured by fitting variogram models to the residuals of the GLM. In addition, we analyzed the species‐area relationship (area=tree diameter). Different environmental variables explained the richness of different species groups (bryophytes vs lichens, specialists vs generalists, sexual vs asexual dispersal). In most groups, the total variation explained by environmental variables was higher than the variation explained by the spatial model. Spatial aggregation was more pronounced in asexually than in sexually dispersed species. Bryophyte species richness was only poorly predicted by area, and lichen species richness was not explained by area at all. Spatial aggregation may indicate effects of dispersal limitation and metapopulation dynamics on community species richness. Our results suggest that species groups differ in habitat requirements and dispersal abilities; there were indications that presence of species with different dispersal strategies is linked to the age of the host tree. Separate analyses of the species richness of species groups that differ in the degree of habitat specialization and dispersal ability give insights into the processes determining community species richness. The poor species‐area relationship, especially in lichens, may indicate species turnover rather than accumulation during the lifetime of the host tree. Epiphyte species extinctions may be mainly caused by deterministic processes, e.g. changes in habitat conditions as the host tree grows, ages and dies, rather than by stochastic population processes.