Ordination of epiphytic bryophyte communities in a wet-temperate coniferous forest,South-Coastal British Columbia

Three coniferous tree species (Pseudotsuga menziesii, Tsuga heterophylla, and Thuja plicata) at Capilano Canyon, British Columbia, were studied for their epiphytic communities. Quantitative data were obtained for fourteen bryophyte species by sampling at heights of 0.5 m, 1 m and 2 m above ground level. Scapania bolanderi is an abundant and widely distributed species; Dicranum fuscescens, Bazzania denudata, Lepidozia reptans and Hypnum circinale are also common. An environmentally uniform study area was chosen to minimize the effects of factors other than bark and elevation on the distribution of epiphytes. In general, tree species are similar in terms of their epiphytic assemblages, but ordination methods using quantitative data exposed compositional variation that may be explained by differences in microclimate and bark-type. In this respect, the results point the way to further studies to examine epiphyte associations in closer detail, and to relate these associations to specific factors in the microenvironment.Nomenclature for vascular plants follows Hitchcock & Cronquist (1973); bryophyte nomenclature follows Schofield (1976) for mosses and Stotler & Crandall-Stotler (1977) for hepatics.We thank Dr. W. B. Schofield for aiding in bryophyte identification.     

Tree Species Composition Predicts Epiphytic Lichen Communities in an African Montane Rain Forest

The ecology of many tropical rain forest organisms, not the least in Africa, remains poorly understood. Here, we present a detailed ecological study of epiphytic lichens in the equatorial montane rain forest of Bwindi National Park (331 km²), Uganda. We evaluated all major lichen growth forms, including selected groups of crustose lichens. In 14 transects at elevations of 1290 m to 2500 m, we sampled 276 trees belonging to 60 species. We recorded all lichen species on each tree trunk between ground level and 2 m above the ground, yielding 191 lichen species in 67 genera, with a mean of 4.7 species per tree. We used non‐metric multi‐dimensional scaling to separate epiphytic lichen assemblages according to tree species composition and elevation. Structural equation modeling indicated that elevation influenced tree species composition and that tree species composition largely determined lichen species composition. Thus, elevation acted indirectly on the lichen assemblages. Further studies examining factors such as bark properties and lichen colonization ecology may clarify what determines the association between tree species and lichen assemblages. The link between lichen assemblages and large‐scale elevation patterns, as well as disturbance and regrowth histories, warrants further study. An analysis of lichen species composition on individual tree species that occur over large elevation ranges would distinguish the effect of tree species on lichen assemblages from the effect of elevation and thus climate. Our study highlights the limited extent of our knowledge of tropical epiphytic lichens.     

云南哀牢山原生林及次生林群落附生地衣物种多样性与分布

附生地衣是森林附生植物的重要类群之一, 在维护森林生态系统的物种多样性以及水分和养分循环等方面发挥着重要作用。作者于2005年12月至2006年5月利用树干取样法调查了云南哀牢山徐家坝地区原生山地常绿阔叶林及其次生群落栎类萌生林、滇山杨(Populus bonatii)林和花椒(Zanthoxylum bungeanum)人工林中525株不同种类和径级树木距地面 0–2.0 m处附生地衣的组成和分布, 并收集了各个群落地面上凋落的地衣, 分析了林冠层附生地衣的物种组成。研究结果表明, 该区森林中附生地衣物种比较丰富。共收集到附生地衣61种, 分属17科29属, 其中原生林、栎类萌生林、滇山杨林和花椒人工林分别有51、53、46和23种。在树干距地面 0–2.0 m位置, 各群落中的附生地衣组成明显不同;但在林冠层中, 各群落内的附生地衣基本相似。原生林中附生地衣种类较多, 但分布不均匀。树干附生地衣的Shannon-Wiener和Simpson多样性指数以栎类萌生林最高, 分别为2.71和0.89;花椒林和滇山杨林次之, 分别为2.43–2.45和0.88–0.89;原生林最低, 为1.25和0.67。树干方位、宿主种类和宿主径级等都对附生地衣的物种组成和多样性有着重要影响, 附生地衣更多地出现于树干南向方位, 云南越桔(Vaccinium duclouxii)的附生地衣最为丰富, 胸径5.0–25.0 cm的树木上附生地衣较多。哀牢山山地森林群落中丰富的附生地衣种类及物种多样性在维系本区山地森林生态系统生物多样性格局方面具有重要的作用。     

Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens. Nomenclature Randlane et al. (2007) for lichens; Leht (2007) for vascular plants.     

Tree hollows can affect epiphyte species composition

Tree hollows often harbor animals and microorganisms, thereby storing nutritive resources derived from their biological activities. The outflows from tree hollows can create unique microenvironments, which may affect communities of epiphytic organisms on trunk surfaces below the hollows. In this study, we tested whether the species richness and composition of epiphytic bryophytes (liverworts and mosses) and lichens differ above and below tree hollows of Aria japonica and Cercidiphyllum japonicum in a Japanese temperate forest. The species richness of epiphytic bryophytes and lichens did not differ above and below hollows; however, the species composition of bryophytes differed significantly above and below hollows. Indicator species analyses showed that the moss species Anomodon tristis and the liverwort species Porella vernicosa were significantly more common below than above hollows, while the liverwort species Radula japonica and four lichen species, including Leptogium cyanescens, occurred more frequently above than below hollows. Our results highlight that tree hollows can produce unique microenvironments on trunk surfaces that potentially contribute to the maintenance of epiphytic diversity on a local scale.     

Correlations among vegetation strata and environmental variables in subalpine spruce-fir forests,southeastern British Columbia

A set of forest vegetation-environment data consisting of 110 plots from subalpine spruce-fir forests in southeastern British Columbia was analyzed with multivariate methods. The aims were to describe and compare relationships among plots using species composition data from four strata of the forest (trees, shrubs, herbs, bryophytes), and to compare patterns in the separate strata with patterns in the environmental data. Weak but statistically significant correlations were detected among PCA axes from all four strata, with a tendency for correlations to be stronger between axes of adjacent strata. Relationships between stratal patterns and environmental variation were examined by correlating PCA axes with environmental variables separately, and jointly with canonical correlation analysis. Linear composites of the environmental data identified by the first canonical axis were similar for shrub, herb, and bryophyte strata implying similar responses to environmental variation in understorey strata; however, < 16% of the total variance was explained. No clear relationships emerged between tree stratum and environmental variation. Probable determinants of pattern in the tree stratum include historical events such as fire and winter snowpack, as well as possible interactions between species. Collectively, these events bear little resemblance to existing environmental conditions in the forest.Nomenclature used by BCMF follows Crum, Steere & Anderson (1973), Hale & Culberson (1970), and Stottler & Crandall-Stottler (1977) for non-vascular plants, and Taylor & MacBryde (1977) for vascular plants.The British Columbia Ministry of Forests provided the data on which this paper is based, as well as a grant to pay for the computer time used. We are grateful to G. Utzig for initially suggesting that a project of this nature be done, and to S. Phelps for helping with the retrieval of computer data files from tape. This paper has greatly benefited from discussions with P. Comeau and R. K. Scagel, and the comments of two anonymous reviewers.     

Leaf demography and leaf traits of temperate-deciduous and tropical evergreen-broadleaved trees in a Mexican montane cloud forest

The spatial variation in epilithic lichen community structure was investigated as part of a larger study of the vegetation and ecology of the tall limestone cliffs within the Niagara Escarpment Biosphere Reserve in southern Ontario, Canada. The cover of all lichen taxa was visually estimated for a total of 199 samples taken from the top, middle, or bottom of the cliff face at five sites. Twelve environmental variables were also measured. Twenty-seven lichen taxa were identified on the samples. Multivariate ordinations of species composition (DCA, CCA, PCCA) revealed variation in community structure on multiple scales, but no groupings of sites that would have suggested the presence of several distinct species assemblages. A gradient in species composition from north to south, most clearly reflected in the decreasing cover of foliose and fruticose species, may reflect a gradient in human disturbance. There was also intermediate-scale patchiness in species composition in a horizontal plane across cliffs, but despite earlier claims made in the literature, no evidence of vertical zonation of the lichens on cliffs was found. Species composition also responded to small-scale factors possibly related to exposure, light, or moisture. Unlike community composition, the total cover of all lichens was homogeneous over large spatial scales and varied only on a small scale, illustrating that scale as well as resolution of a study may influence the ecological patterns seen. More than half of the species found on the Niagara Escarpment are rare on rock substrates elsewhere in southern Ontario, and two are new for North America (Candelariella heidelbergensis (Nyl.) Poelt and Lecanora perpruinosa Fröberg). The result that cliffs support a distinct flora containing many rare species suggests that they are a reservoir for biodiversity not just for vascular plants, but also for lichens.     

DISTRIBUTION AND SUCCESSION OF LICHENS ASSOCIATED WITHPRUNUS SPINOSAAT DUNGENESS,ENGLAND

The distribution and succession of epiphytic and other associated lichens on blackthorn (Prunus spinosa) growing on the shingle at Dungeness is described. Some 33 epiphytic species, which exhibited a range of ecological strategies from pioneer to late colonizers, were recorded. Both the rate of change of status (age) of blackthorns and the progress of lichen colonization seemed to depend on whether the bushes were growing directly on shingle or on adjacent alluvial soil; both processes seeming to be more rapid on the former substratum. Terricolous lichens, of which 18 were recorded, generally appeared late in the succession and mostly amongst bushes growing on shingle. Rabbits often invaded the blackthorns, especially those on alluvium, and probably contributed to the lack of terrestrial lichens under these bushes. Blackthorns growing on shingle tended to be more prostrate in form than those on alluvium. 1996 The British Lichen Society     

Spatial distributional patterns of macroinvertebrates along rivers within and among biomes

This study was designed to test the biome dependency hypothesis, which predicts that similar assemblages of macroinvertebrates occur along rivers both within and among drainage basins if the basins occupy the same biome. Benthic macroinvertebrates were collected from three drainage basins within each of three biomes in Canada, the eastern deciduous forests (EDF) of southwestern Ontario, the grasslands of south-central Alberta, and the montane coniferous forests (MCF) of southeastern British Columbia. A total of 225 benthic samples (3 biomes × 3 rivers/biome × 5 sites/river × 5 samples/site) was collected in spring using a cylinder sampler.The significant interaction effect between biome and a site's location along a river indicated that spatial patterns of variation in total density and taxonomic composition were not spatially consistent among sites along rivers or among biomes. Total macroinvertebrate densities were equivalent between the EDF and grassland sites. However, total density was substantially lower at the MCF sites than at sites in the other two biomes. The greatest differences in taxonomic composition occurred among biomes, although significant differences also occurred for all other sources of variation examined. Macroinvertebrate composition was more strongly associated with local, site-specific factors (riparian vegetation and land use) than with longitudinal gradients. Distinct site-specific taxonomic assemblages were evident in EDF, but not in the other two biomes where land use was more homogeneous.     

Understorey vegetation in boreal Picea mariana and Populus tremuloides stands in British Columbia

Abstract. We compared the species composition and species density of vascular plants in the understorey vegetation of boreal forest between Picea mariana (Black spruce) and Populus tremuloides (Trembling aspen) stands in British Columbia, Canada, and related differences in species composition and species density between the two forest types to dominant canopy tree species as well as a wide variety of environmental factors. We analysed 231 stands, distributed in three different climatic regions representing drier, wetter, and milder variations of montane boreal climate. Of these stands 118 were dominated by P. mariana and 113 by P. tremuloides. P. tremuloides stands had higher species density than P. mariana stands in all climatic regions, but species density of each dominance type varied among climatic regions. The floristic composition of the understorey vegetation was markedly different for P. mariana and P. tremuloides dominated stands. A detailed study on the effect of canopy dominance and local environmental factors on the understorey vegetation of the boreal forest was conducted using 88 stands from one of the three climatic regions. Using a combination of ordination and variation partitioning by constrained ordination we demonstrated a small but unique effect of canopy dominance type on the understorey vegetation, while a larger amount of compositional variation was shared with other factors. Our results accord with a scenario in which differences in primary environmental factors and humus form properties, the latter accentuated by the canopy dominants themselves, are the most important causes of higher species density in P. tremuloides stands than in P. mariana stands, as well as differences in species composition among the two canopy dominance types. Processes and time scales involved in the small but significant direct and indirect effects of the canopy dominant on understo‐ rey species composition are discussed.     

Analysis of the distribution of epiphytic lichens within homogeneousFagus sylvatica stands along an altitudinal gradient (Mount Olympos,Greece)

Epiphytic lichen vegetation onFagus sylvatica sas studied in 4 sites along an altitudinal gradient from 930 to 1500 m on SE facing slopes of Mount Olympos (Greece). The crucial factor determining the spatial heterogeneity of epiphytic lichens onF. sylvatica is the altitude and not the height on the trunk at which lichen community is established. 17 out of 26 taxa are confined to a particular elevation range, while another three are clearly ubiquitous in their distribution. The number of lichen species at breast height is higher than at the base of the trunks. The results were compared with those gathered earlier in an analogous study on the vertical distribution of epiphytic lichens onPinus nigra along an altitudinal gradient from 750 to 1510 m of the same mountain. Comparison suggests that spatial heterogeneity of epiphytic lichens onF. sylvatica is different from the one onP. nigra.     

Epiphytic lichen synusiae and functional trait groups in boreo‐nemoral deciduous forests are influenced by host tree and environmental factors

Deciduous forests with temperate broad‐leaved tree species are particularily important in terms of biodiversity and its protection, but are threatened habitats in northern Europe. Using multivariate analyses we studied the effect of forest site type, environmental variables and host tree properties on epiphytic lichen synusiae as well as on the composition of species‐specific functional traits. Epiphytic lichens were examined on Acer platanoides, Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra and U. laevis in two types of forests: Humulus‐type floodplain forests and Lunaria‐type boreo‐nemoral forests on the talus slopes of limestone escarpment (klint forests). Klint forests located near the seashore were under greater maritime influence compared to floodplain forests located in inland Estonia which experience stronger air temperature contrasts. In addition to stand level and climatic variables, tree level factors (bark pH, trunk circumference and cover of bryophytes) considerably affected the species composition of the lichen synusiae. Overall, 137 lichen species were recorded, including 14 red‐listed species characteristic of deciduous trees. We defined 13 lichen societies and showed their preference to forests of a specific site type and/or host tree properties. In forests of both types, most of the epiphytic lichens were crustose, and had apothecia as the fruit bodies and chlorococcoid algae as the photobiont. However, the proportion of lichens with a foliose or fruticose growth form, as well as the proportion of lichens with vegatative diaspores, were higher in floodplain forests. In klint forests with a stronger influence from the wind, crustose species completely dominated, while species with vegetative diaspores were rare and most species dispersed sexually. Lichens with Trentepohlia as the photobiont were characteristic of these forests, and lichens with lirellate ascomata were prevailing, indicating the great uniqueness of the kint forests for epiphytic lichens in the boreo‐nemoral region.     

Effects of epiphytic lichens on host preference of the vascular epiphyte Tillandsia usneoides

We investigated the potential for nonvascular epiphytic species (primarily lichens) to affect the quality of different host tree species for the vascular epiphyte Tillandsia usneoides in the southeastern USA. Different host tree species had substantially different abundances of Tillandsia , and these abundances were correlated with the composition of nonvascular epiphyte communities. In greenhouse experiments Tillandsia grew significantly faster on the branches of Quercus virginiana (a species with very high natural abundances of Tillandsia ) when the dominant lichen on Q. virginiana was left intact than when the lichen was removed from the branches. In laboratory experiments, extracts from Cryptothecia rubrocincta , a lichen that was 10 times more common on poor host species for Tillandsia than on good host species, reduced Tillandsia seedling survival and growth in comparison to extracts from other species and rainwater. In field experiments, lichens increased the proportion of Tillandsia seeds and vegetative strands that adhered to the trunk of Ilex opaca (a poor Tillandsia host), but lichens did not affect propagule adherence to Q. virginiana . Our results are by no means exhaustive of the possibilities, but they suggest that the structure and diversity of vascular and nonvascular epiphytic communities that grow in different tree species may not be simply the product of host tree characteristics, but may also be influenced by interactions among the epiphytes themselves.     

Epiphytic and epixylic lichen species diversity in Estonian natural forests

The epiphytic and epixylic lichen flora of natural forests was recorded in different parts of Estonia. Altogether 232 taxa of lichens, lichenicolous fungi, or non-lichenized fungi were recorded, 10 of them listed in the Estonian Red Data Book. We found regional differences in lichen species composition and diversity caused by differences in the forest types. The tree-species-rich boreo-nemoral forests had the most diverse lichen flora, while the boreal forest dominated by coniferous trees or birch had the lowest diversity. The stand age proved to be significant in regard to the number of lichen species in a forest. The most remarkable effect on the diversity of forest lichen species was caused by the presence of Populus tremula. Aspen had the highest number of lichen species on the basal trunk and twigs, and also the highest number of host-specific lichen species.     

Assessing the effects of forest management on epiphytic lichens in coppiced forests using different indicators

Abstract

Epiphytic lichens are one of the taxonomic groups most sensitive to forest management. Nevertheless, they have not yet been exhaustively included in the assessment of Sustainable Forest Management. This work aimed at evaluating the effects of forest management on epiphytic lichens in coppiced forests, exploring the spatial patterns of diversity and the composition of communities. Moreover, the goal was to compare the performance of four potential indicators for monitoring the effects of forest management on epiphytic lichens: total lichen diversity, species associated with intensive management, species associated with aged coppiced woodlands and Indicator Species Ratio (ISR). In humid Mediterranean Liguria, 50 sampling units were chosen in Castanea sativa and deciduous Quercus spp. forests subjected to different forest management practices: intensively managed coppice and aged coppice/high forest. The effect of forest management was evident in terms of species composition, since it was possible to find significantly associated species for each of the two management types. At each sampling site, the four indicators were calculated using Indicator Value Analysis and compared through correspondence analysis. The ISR was shown to be a more effective indicator, being independent of floristic composition and the occurrence of rare species.     

A quantitative study of the epiphytic vegetation on pollarded trunks of Fraxinus excelsior at Havrå, Osterøy,western Norway

The epiphytic vegetation on 19 pollarded trees of Fraxinus excelsior at the farm Havrå, Osterøy, western Norway was investigated. Each trunk was divided into a basal zone, a middle zone and a top zone. In each zone the four different aspects were analysed by squares (12 sampling units from each trunk). Within a total of 225 sampling units, 173 taxa were recorded (84 lichens, 72 bryophytes, 17 vascular plants). The epiphytic vegetation was classified into seven TWINSPAN groups, and it is shown by detrended correspondence analysis (DCA) that these groups are mainly separated by the first two DCA axes. In an attempt to explain the floristic composition, several environmental variables were measured, and the floristic and environmental data were analysed by canonical correspondence analysis (CCA). A pollarded tree, depending on its age, represents a continuum of perhaps several hundred years. Some of the recorded epiphytic lichens are very rare in Norway, and the pollarded trees thus contribute greatly to the biodiversity of the old cultural landscape. Several epiphytic species take advantage of favourable light conditions on trunks in the open fields. At Havrå, woodland has developed by tree colonization of old meadows and open fields during the last 40 years after the cessation of earlier managements. Some floristic differences between the epiphytic vegetation on trunks in the open fields and in the woodland are found. This suggests a change in the epiphytic vegetation because of the creation of a more shady habitat in parts of the area during a few decades.     

Patterns and drivers of lichen species composition in a NW-European lowland deciduous woodland complex

We surveyed lichens in an extensive area of lowland deciduous oak and beech dominated forest under wood pasture management within the New Forest National Park, southern England. This provided the basis for an investigation of factors affecting the species density and composition of epiphytic lichen communities. Fifteen 1 hectare plots were established in the same sites as a parallel invertebrate survey, of which nine were in old growth forest and six in oak plantations of between c. 150 and 300 years old. In each plot 12 trees were identified for sampling and lichens were sampled on four aspects of each trunk. Results at the plot level showed that species density was significantly higher in the old growth woodland plots from those in plantations. Plot age had a significant effect on species on oak but this was not significant for lichen communities on beech. The species density of lichens associated with Trentepohliaceae photobionts on oaks showed a linear positive relationship with increasing plot age while the species density of species with other coccal green Chlorophyta decreased with age of the plot. A major part of the lichenised fungi with Trentepohliaceae photobionts includes indicator and notable species used in conservation evaluation and this component is most affected by fragmentation and isolation of forest sites. The results emphasise the importance of long term connectivity and ecological continuity in this extensive mosaic of lowland deciduous forest.     

Variation in soil seed bank species composition of a dry coniferous forest: spatial scale and sampling considerations

In order to assess the effects of disturbance on soil seed bank spatial structure, variation in seed bank species composition was analyzed at two spatial scales in mature (undisturbed) and recently disturbed stands of Interior Douglas-fir forest in south-central British Columbia. Coarse-scale analysis among stands 10–100 km apart using Multi-Response Permutation Procedures showed that individual mature stands were mainly characterized by unique seed bank species compositions. Disturbance had an homogenizing effect on seed bank species composition at the coarse scale––there was less variability among stands following disturbance by both low and high severity fires, and by non-salvage logging. In contrast, finer scale analysis among sampling units c. 10 m apart showed that heterogeneity in seed bank species composition was greater on severely burned and logged sites, commensurate with greater levels of soil disturbance, than on lightly burned and undisturbed sites. Despite the high intensity of seed bank sampling (sixty 25 cm² soil samples from each of 16 sites), species-area curves leveled off only when infrequent species were removed from the dataset. The number of seed bank samples required to account for the common species ranged from 84 on severely burned sites, to 196 on undisturbed sites; more than the 240 samples collected per disturbance class in this study would have been required to account for the remaining infrequent species. Overall, this study highlights the importance of sampling intensively within multiple stands to capture the variation in species composition inherent to these dry coniferous forest soil seed banks.     

Microsatellites reveal regional population differentiation and isolation in Lobaria pulmonaria, an epiphytic lichen

Many lichen species produce both sexual and asexual propagules, but, aside from being minute, these diaspores lack special adaptations for long-distance dispersal. So far, molecular studies have not directly addressed isolation and genetic differentiation of lichen populations, both being affected by gene flow, at a regional scale. We used six mycobiont-specific microsatellite loci to investigate the population genetic structure of the epiphytic lichen Lobaria pulmonaria in two regions that strongly differed with respect to anthropogenic impact. In British Columbia, L. pulmonaria grows in continuous old-growth forests, while its populations in the old cultural landscape of Switzerland are comparably small and fragmented. Populations from both British Columbia and Switzerland were genetically diverse at the loci. Geographically restricted alleles, low historical gene flow, and analyses of genetic distance (upgma tree) and of differentiation (amova) indicated that populations from Vancouver Island and from the Canadian mainland were separated from each other, except for one, geographically intermediate population. This differentiation was attributed to different glacial and postglacial histories of coastal and inland populations in British Columbia. In contrast to expectations, the three investigated Swiss populations were genetically neither isolated nor differentiated from each other despite the long-lasting negative human impact on the lichen's range size in Central Europe. We propose that detailed studies integrating local landscape and regional scales are now needed to understand the processes of dispersal and gene flow in lichens.