Local extent of old-growth woodland modifies epiphyte response to climate change

Aim To quantify the interaction between climate and woodland continuity in determining the bioclimatic response of lichen epiphytes. Location Northern Britain (Scotland). Methods Indicator‐species analysis was used to pre‐select lichen epiphytes along parallel gradients in climate and the extent of old‐growth woodland. Nonparametric multiplicative regression was used to describe in a predictive model the individualistic response of selected species, which were projected based on climate‐change scenarios and contrasting patterns of simulated woodland loss or gain. Species with a similar response were grouped using a novel application of cluster analysis to summarize the potentially huge number of projected outcomes. Projected patterns of occurrence under climate‐change scenarios were examined for different levels of old‐growth woodland extent. Results Forty‐two lichen species were statistically significant indicator species in oceanic woodlands, and old‐growth indicators under suboptimal climatic conditions. Responses to climate‐change scenarios were contrasting, with one group comprising species projected to increase in extent in response to climate warming, and other response groups projected to decrease in occurrence, possibly in response to shifting rainfall patterns. The occurrence of all response groups had a positive relationship with old‐growth woodland extent. Main conclusions An ‘oceanic’ biogeographical group of epiphytes identified using the baseline climatic and present‐day woodland setting comprised species with a cyanobacterial photobiont or tropical phytogeographical affinities. However, within this group the individual species responses to climate‐change scenarios were contrasting. Additionally, group responses may be poorly matched with simple ecological traits. However, the studied interaction between climate and habitat continuity suggests that the impact of climate change might be offset for certain lichen epiphytes by appropriate management of woodland resources, for example, expansion of native woodland around remnant old‐growth stands.     

Report of the Annual Conference of the Cryptogamic Section, 1948

Background: Trait-based assembly rules are a powerful tool in community ecology, used to explore the pattern and process of community structure (richness and composition).

Aims: A preliminary test for the utility of trait-based assembly rules in explaining cryptogamic epiphyte communities (lichens and bryophytes).

Methods: We sampled epiphytes from three different tree species (aspen, birch and pine), and from trees of contrasting age. The community composition of epiphyte species (taxon analysis) and functional groups (trait analysis) was summarised using multivariate ordination (nonmetric multidimensional scaling, NMDS).

Results: Ordination documented a widely observed pattern in which different tree species have taxonomically different epiphyte communities. However, NMDS sample scores were correlated to tree age in the trait-based analysis, but not in the taxon analysis.

Conclusions: Our results point to the existence of a common pattern in community traits during succession (on trees of different age) when measured for epiphyte communities with contrasting taxonomic composition. This pattern is evidenced by consistent trends in lichen growth form and reproductive strategy (sexual vs. asexual).     

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.     

Lichen epiphyte diversity: A species,community and trait-based review

The forest canopy is fundamentally important in biodiversity conservation and ecosystem function. Cryptogamic epiphytes are dominant tree bole and canopy elements in temperate and boreal forests, though remain neglected by mainstream forest ecology. This review makes ecological information on cryptogamic epiphytes available to a non-specialist audience, to facilitate their integration in forest biodiversity and ecosystem studies more generally. The review focuses specifically on lichen epiphytes, highlighting their diversity and ecosystem role. A principal task is to explore pattern and process in lichen epiphyte diversity – species composition and richness – therefore demonstrating the utility of lichens as an ecological model system. The review examines key themes in previous research. First, the extensive literature used to resolve species response to, and community turnover along environmental/resource gradients, consistent with the habitat niche. Second, the evidence for dispersal-limitation, which may constrain community composition and richness in isolated habitats. Third, these two processes – the habitat niche and dispersal-limitation – are used to explain stand-scale diversity, in addition to the role of neutral effects (habitat area). Fourth, the review moves from a taxonomic (pattern) to a functional (process) perspective, considering evidence for autogenic succession evidenced by competition and/or facilitation, and non-random trends in life-history traits. This functional approach provides a counter-point to an assumption that lichen epiphyte communities are unsaturated and non-competitive, a situation which would allow the long-term accumulation of species richness with temporal continuity. Finally, the review explores landscape-scale impacts on lichen epiphytes, with recommendations for conservation.     

Archaeobotanical evidence for climate as a driver of ecological community change across the anthropocene boundary

The biodiversity response to climate change is a major focus in conservation research and policy. Predictive models that are used to project the impact of climate change scenarios – such as bioclimatic envelope models – are widely applied and have come under severe scrutiny. Criticisms of such models have focussed on at least two problems. First, there is an assumption that climate is the primary driver of observed species distributions (‘climatic equilibrium’), when other biogeographical controls are often reliably established. Second, a species' sensitivity to macroclimate may become less relevant when impacts are down‐scaled to a local level, incorporating a modifying effect of species interactions structuring communities. This article examines the role of different drivers (climate, pollution and landscape habitat structure) in explaining spatial community variation for a widely applied bioindicator group: lichen epiphytes. To provide an analysis free of ‘legacy effects’ (e.g. formerly higher pollution loads), the study focused on hazel stems as a relatively short‐lived and recently colonized substratum. For communities during the present day, climate is shown to interact with stem size/age as the most likely explanation of community composition, thus coupling a macroclimatic and community‐scale effect. The position of present‐day communities was projected into ordination space for eight sites in England and compared to the position of historical epiphyte communities from the same sites, reconstructed using preserved hazel wattles dating mainly to the 16th Century. This comparison of community structure for the late‐ to post‐Mediaeval period, with the post‐Industrial period, demonstrated a consistent shift among independent sites towards warmer and drier conditions, concurrent with the end of the Little Ice Age. Long‐term temporal sensitivity of epiphyte communities to climate variation thus complements spatial community patterns. If more widely applied, preserved lichen epiphytes have potential to generate new baseline conditions of environment and biodiversity for preindustrial lowland Europe.     

Integrating multiple landscape-scale drivers in the lichen epiphyte response: climatic setting, pollution regime and woodland spatial-temporal structure

Aim To quantify the role of multiple biodiversity drivers – pollution, woodland structure and climate – controlling lichen epiphyte composition and diversity. Location  Scotland, north‐west Europe. Methods Four compatible datasets were assembled: site‐scale species distribution data (response) and base‐line modelled data on climate, pollution loads and extent of old‐growth woodland (explanatory variables). First, partial‐canonical correspondence analysis was used: (1) to compare the importance of environmental variables to pure spatial effects and (2) to partition the importance of environmental variables in explaining species composition. Secondly, patterns of species richness were investigated using multiple least‐squares regression. Results Old‐growth woodland was the most important control of species richness. Pollution was the most important explanatory variable for species composition. The impact of pollution on composition (and to a lesser extent on richness) is explained: (1) By recovery of lichens with declining SO₂ pollution, although with epiphyte composition shifted by the recent effects of N‐pollution and (2) By the limited spatial extent of severe pollution, and generally low‐to‐moderate pollution loads across our study area, combined with the positive effect of old‐growth woodland extent in controlling species richness. The effect of climate and old‐growth woodland on species composition covaried, supporting an interaction between habitat quality and climatic setting, which may be important in understanding the epiphyte response to climate change. Conclusions Advances in conservation planning will likely require an integrated approach to understanding simultaneous effects of multiple drivers, providing opportunities for integrated management strategies. Our study provides a preliminary example of this approach by combining three key biodiversity drivers into a single framework for lichen epiphytes. Thus, reducing pollution loads may make old‐growth woodland that currently exists in a polluted landscape available for colonization, thereby extending the available habitat for epiphytes, and facilitating an effective species response to climate change.     

Changing climate and historic‐woodland structure interact to control species diversity of the ‘Lobarion’ epiphyte community in Scotland

Question: How will changing climate and habitat structure interact to control the species diversity of lichen epiphytes? Location: Scotland. Method: Species richness (=diversity) of the epiphyte lichen community known as Lobarion (named after Lobaria pulmonaria) was quantified for 94 Populus tremula stands across Scotland, and compared in a predictive model to seven climate variables and eight measures of woodland structure. An optimum model was selected and used to project Lobarion diversity over the geographic range of the study area, based on IPCC climate change scenarios and hypothetical shifts in woodland structure. Results: Species diversity of the Lobarion community was best explained by three climate variables: (1) average annual temperature; (2) autumn and winter precipitation; in combination with (3) historic‐woodland extent. Projections indicate a positive effect of predicted climate change on Lobarion diversity, consistent with the physiological traits of cyanobac‐terial lichens comprising the Lobarion. However, the general response to climate is modified significantly by the effect on diversity of historic‐woodland extent. Conclusions: Historic‐woodland extent may exert an important control over local climate, as well as impacting upon the metapopulation dynamics of species in the Lobarion. In particular, a temporal delay in the response of Lobarion species to changed woodland structure is critical to our understanding of future climate change effects. Future Lobarion diversity (e.g. in the 2050s) may depend upon the interaction of contemporary climate (e.g. 2050s climate) and historic habitat structure (e.g. 1950s woodland extent). This is supported by previous observations for an extinction debt amongst lichen epiphytes, but suggests an extension of simple climate‐response models is necessary, before their wider application to lichen epiphyte diversity.     

The impacts of beavers Castor spp. on biodiversity and the ecological basis for their reintroduction to Scotland,UK

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Diversity and succession of epiphytic macrolichen communities in low‐elevation managed conifer forests in Western Oregon

Abstract. We examined epiphytic macrolichen communities in Pseudotsuga menziesii (Douglas‐fir) forests across the western Oregon landscape for relationships to environmental gradients, stand age and structure, and commercial thinning. We used a retrospective, blocked design through the Coast and the western Cascade ranges of Oregon. Each of our 17 blocks consisted of a young, unthinned stand (age 50–110 yr); an adjacent, thinned stand of equivalent age; and an old‐growth stand (age > 200 yr). We found 110 epiphytic macrolichen taxa in the stands. Forage‐providing alectorioid lichens and the nitrogen‐fixing cyanolichen Lobaria oregana associated strongly with old‐growth stands and remnant old trees in younger stands (unthinned + thinned). Relative to unthinned stands, thinned stands had a slightly higher abundance of alectorioid lichens and a greater presence of Hypogymnia imshaugii. However, thinned stands hosted a lower landscape‐level (γ) diversity, lacking many species that occurred infrequently in the unthinned stands. Patterns in the lichen community composition correlated strongly with climatic gradients; the greatest variation in composition was between the Coast and Cascade ranges. The difference in communities between mountain ranges was greatest among stands 70–110 yr old, suggesting a difference in lichen successional dynamics between the ranges.     

The potential contribution of conifer plantations to the UK Biodiversity Action Plan

Summary

The detrimental effects of conifer plantations on open ground habitats have been well catalogued and discussed, but the potential contribution of planted forests to the conservation of woodland biodiversity has not been quantified to the same extent. This quantification is needed urgently to help forest managers fulfil commitments to biodiversity enhancement as outlined in the UK Biodiversity Action Plan, the UK Forestry Standard and the UK Woodland Assurance Scheme (UKWAS). Results are presented from a five-year programme of research aimed at obtaining baseline information on biodiversity in planted forests and evaluating the contribution of planted forests to the conservation of native flora and fauna. Fifty-two plots were surveyed in total, covering a range of different tree crops (Scots pine Pinus sylvestris L., Sitka spruce Picea sitchensis (Bong.) Carr., Norway spruce Picea abies L. and Corsican pine Pinus nigra var maritima (Aitón) Melville) and stand ages (pre-thicket, mid-rotation, mature and over-mature) in three contrasting bioclimatic zones (upland, foothills and lowlands) throughout Britain. Additional plots were established in semi-natural woodland to allow comparisons between the biodiversity of plantations and native stands. Over 2000 species were recorded in total, including 45 Red Data Book species. Planted stands had similar or richer fungal and invertebrate communities to those of the native stands but poorer lichen and vascular plant communities. The latter were strongly affected by shading, dense, mid-rotation Sitka spruce stands having the lowest species counts. In contrast, these stands had a high diversity of mycorrhizal fungi, including a number of rare and threatened species normally associated with native pine wood. Bryophyte species-richness was related more to climate than woodland type, with the wetter upland spruce and native oak stands having the most diverse communities. Compared to the younger planted stands, over-mature planted stands had a higher proportion of species characteristic of semi-natural woodland stands. This related to greater structural diversity and higher deadwood volumes in the over-mature stands. It is concluded that conifer plantations make a positive contribution to biodiversity conservation in the UK and hence to the UK Biodiversity Action Plan. No single stand or crop type provides ‘optimal’ conditions for biodiversity, but the habitat value of plantations could be enhanced by increasing the area managed under alternative systems to clear-felling, such as ‘continuous cover’ and/or non-intervention natural reserves.     

Lichens — the biodiversity value of western woodlands

Summary

The Atlantic broad-leaved woodlands of Britain are of international renown for their lichen floras. They are inhabited by 517 lichens, representing 28.3% of the total lichen flora and 73.2% of all British woodland lichens, and they are the main habitat for 165 species. Of these, 31 have a marked southern distribution and do not reach Scotland, whereas 26 species are found in Scotland, but not England or Wales. Their British Red-listed species are outnumbered by the 86 species for which Britain has International Responsibility.

Within the Atlantic broad-leaved woodlands, only 30 lichens show a preponderance for oak. With the exception of some ancient oakwoods in southern England, a high lichen biodiversity is rarely dependent on a dominance of oak in the woodland canopy, more usually it is the result of a long ecological continuity, often a varied tree and shrub composition, a varied canopy density, and good air quality. Consequently, the oak stands within former ‘industrial’ woodlands have a much lower lichen biodiversity compared with woodlands that have a history as ‘pasture woodland’ or, as with some ravine woodlands, have otherwise escaped intensive management.

The life-history of an oak tree is considered in relation to the niches it provides for lichen colonisation with time. Some management scenarios are provided with the enhancement of the lichen interest of former ‘industrial’ oakwoods as an objective.     

Epiphytic (including hemiepiphytes) diversity in three timber species in the southwestern Amazon, Brazil

Forestry managers have been searching for ways to reduce the impacts of logging on Amazonian biodiversity, but some basic factors are still not considered in native forestry operations, among them the diversity of epiphytes associated with the logged trees. Our goals in this study were to determine the floristic composition, quantify the species richness, and characterize the species diversity of the vascular epiphytic community present in three timber tree species in Acre State, Brazil. We collected and identified all epiphytes in 30 randomly selected trees ≥35 cm DBH of each of three important timber species, Tabebuia serratifolia, Manilkara inundata and Couratari macrosperma. We also documented the epiphyte diversity in 120 randomly selected trees ≥35 cm DBH of 56 other species to determine whether the three timber species have different epiphyte diversity than the tree community at large. The epiphyte samples in the three timber species showed 77 species, 13 of which were new records for the flora of Acre state. The epiphyte community in the randomly selected trees presented a total of 56 species. The timber species phorophytes hosted on average three times more epiphyte species per tree than the other 120 randomly selected trees. These results show that a substantial portion of local floristic richness can be lost during logging activity due if not properly managed by rescuing epiphytes after felling the trees. Although these epiphytes could contribute positively to forestry sustainability due to their ornamental value, increasing the economic yield per hectare, there are no local initiatives for economic use of epiphytes.     

Recolonization of vascular epiphytes in a shaded coffee agroecosystem

Aim: Shaded coffee plantations constitute an important refuge for biodiversity. Despite the fact that epiphytic plants form a significant component of these agroecosystems, their removal from the shade trees is commonplace in Latin America. To what extent does the epiphyte community recover from this severe disturbance? Location: Shaded coffee agroecosystem in Veracruz, Mexico (19°28′03″ N, 96°55′58″ W; 1200 m asl). Methods: We assessed the diversity, biomass and recolonization patterns of vascular epiphytes in shade trees, 8‐9 yr after complete epiphyte removal (E^?), and in control ‘non‐removal’ sites (E⁺). In order to evaluate the effects of prior epiphyte removal, all vascular epiphytes were completely removed from 10 trees per treatment (E^? and E⁺); all epiphyte species collected were identified and dry biomass measured. Results: Eight to nine years after removal, epiphyte biomass in the E^? shade trees was 35% of that found in the control sites. A total of 55 epiphyte species, belonging to 12 families, were registered; 40 in E^?, and 48 in E⁺. Six species belonging to Bromeliaceae, Orchidaceae, Cactaceae and Araceae accounted for 75% of the biomass in E⁺ while six species of bromeliads accounted for 76% of the biomass in E^?. Some bromeliads proliferated following disturbance; however, ferns showed lower recovery. Conclusions: Epiphyte community recovery, in terms of biomass and diversity, is considerably higher in the coffee plantation than has been previously reported for other tropical ecosystems. Epiphyte recolonization patterns reflected both the abundance of species in the surrounding matrix and certain species‐specific traits. For such agroecosytems to function as effective reservoirs of epiphyte diversity, epiphyte stripping should be avoided.     

19th century woodland structure controls stand-scale epiphyte diversity in present-day Scotland

Axioms developed from island biogeography theory (i.e. species–area relationships, effects of fragmentation and isolation) are central to the development of conservation strategy. Within this context, the 'extinction debt' hypothesis brings into question an often assumed relationship between species richness and present-day spatial habitat structure (i.e. extent, fragmentation), suggesting instead that the richness and composition of biological communities may lag behind spatial changes in habitat. We examined evidence for an extinction debt among epiphytic lichens, a highly diverse biological group of significant conservation concern. Using sites in Scotland, we compared epiphyte species richness in smaller-scale habitat units (aspen stands) to larger-scale woodland structure (extent and fragmentation) measured at two spatial scales (1 km² and 4 km²) and for two timeframes, modern (1990s to 2000s) and historic (1860s to 1880s). Species richness was positively related to woodland extent and negatively related to woodland fragmentation; however, richness was explained better by historic woodland structure at a 1-km² scale, than by modern woodland structure. The results indicate: (1) a coupling of stand-scale epiphyte assembly and dynamics of the wider woodland ecosystem, and (2) a significant lag in the response of epiphyte species richness to habitat spatial structure. However, the effect of spatial habitat structure is different between species groups with contrasting traits. The effect of decreasing woodland extent on epiphyte richness is generally more severe for microlichens (comprising a greater number of rare and specialist species) than the more generalist macrolichens.     

Epiphyte associations and canopy soil volume: nutrient capital and factors influencing soil retention in the canopy

• Canopy soil (CS) volume reflect epiphyte community maturity, but little is known about the factors that retain CS or species succession within it. Humus fern species (e.g. Phlebodium areolatum) appear capable of retaining CS.
• In ten Quercus spp. we sampled 987 epiphyte mats to examine the role of the common epiphyte species and crown traits determining CS volume, in order to infer successional stages and identify pioneer and late successional species. Branch traits (height, diameter and slope), CS volume and cover of the epiphyte species were determined for each mat. Nutrient content was determined in CS random samples of 12 epiphyte associations and sizes (one sample from each size quintile).
• A total of 60% of the mats lack CS. Cover of P. areolatum was the main variable explaining CS volume, and this species was present in 46.8% of those with CS. Epiphyte composition was highly variable, but pioneer (species appearing in monospecific mats, without CS) and late successional species could be identified. Canopy soil nutrient content was similar among the associations of epiphytes. Magnesium, Ca and pH decreased with CS volume, while P and N increased.
• Phlebodium areolatum is associated with high CS volumes and could act as a key species in its retention. Monospecific mats of pioneer species lack CS or have low volumes, while CS is much higher in mats with late successional species, but the mechanisms of CS formation and nutrient retention in response to interactions between epiphyte species remain to be tested.

     

Can spatial variation in epiphyte diversity and community structure be predicted from sampling vascular epiphytes alone?

Aim Non‐vascular epiphytes have been largely ignored in studies examining the biotic and abiotic determinants of spatial variation in epiphyte diversity. Our aim was to test whether the spatial patterning of species richness, biomass and community composition across geographic regions, among trees within regions, and among branches within trees is consistent between the vascular and non‐vascular components of the temperate rain forest flora. Location Coastal lowland podocarp‐broadleaved forests on the west coast of the South Island of New Zealand. Methods We collected single samples (30 × 25 cm) from 96 epiphyte assemblages located on the inner branches of 40 northern rata (Metrosideros robusta) trees. For each sample, branch characteristics such as branch height, branch diameter, branch angle, branch aspect, and minimum and maximum epiphyte mat depth were recorded. The biomass for each individual epiphyte species was determined. Results Northern rata was host to a total of 157 species, comprising 32 vascular and 125 non‐vascular species, with liverworts representing 41% of all species. Within epiphyte mats, the average total organic biomass of 3.5 kg m^?2 of branch surface area consisted largely of non‐living biomass and roots. Vascular and non‐vascular epiphytes showed strikingly different spatial patterns in species richness, biomass and composition between sites, among trees within sites, and among branches within trees, which could not be explained by the branch structural characteristics we measured. The two plant groups had no significant association in community composition (r = 0.04, P = 0.08). However, the species richness of vascular plant seedlings was strongly linked to the presence/absence of lichens. Main conclusions Non‐vascular plants contributed substantially to the high species richness and biomass recorded in this study, which was comparable to that of some tropical rain forests. High variability in community composition among epiphyte mats, and very low correlation with any of the environmental factors measured possibly indicate high levels of stochasticity in seed or spore colonization, establishment success or community assembly among branches in these canopy communities. Although we found some evidence that vascular plant seedling establishment was linked to the presence of lichens and the biomass of non‐living components in the epiphyte mats, there was no correlation in the spatial patterning or determinants of species richness between non‐vascular and vascular plants. Consequently, variation in total epiphyte biodiversity could not be predicted from the measurement of vascular plant diversity alone, which highlights the crucial importance of sampling non‐vascular plants when undertaking epiphyte community studies.     

Invasion of a Habitat-Forming Seaweed: Effects on Associated Biota

Fucus evanescens is a brown alga of arctic origin that has invaded European coasts. The epiphytic community of F. evanescens in southern Sweden was compared with that of the native Fucus vesiculosus, to examine to what extent an invading seaweed can modify local biodiversity. F. evanescens was much less fouled than F. vesiculosus, supporting both less biomass and fewer species of epiphytes. Multivariate analysis of the most common epiphyte taxa showed that the epiphytic community composition of F. evanescens was not entirely separated from that of F. vesiculosus, but host species contributed significantly to explain the variation in community composition. The biomass of free-living invertebrates was also lower on F. evanescens, although the pattern differed between taxonomic groups. While the biomass of amphipods was lower on F. evanescens, there was no significant difference in biomass of isopods or gastropods between the Fucus species. The good correlation between biomass of epiphytes and free-living animals suggests that the epiphytes play an important role in providing a suitable habitat for many species of free-living epifauna. The study shows that the invasion of F. evanescens affects the environmental conditions for many species associated with the Fucus community but that the direct effect on biodiversity is probably low.     

The epiphyte vegetation of Annona glabra on Barro Colorado Island, Panama

Aim Information on the community composition, structure, and dynamics of epiphyte vegetation is scarce. A survey of the epiphytes occurring on all individuals of one particular host tree species in a well-studied neotropical research site allowed us a comparison of the epiphyte flora of this tree with the local epiphyte flora, the analysis of spatial distribution patterns and the use of these patterns as indications for changes in time. In the future, our results can be used as a baseline data-set for the direct observation of the long-term dynamics in epiphyte communities. Location The study was conducted on Barro Colorado Island (BCI), Panama. Methods We recorded all individuals of the vascular epiphytes growing on Annona glabra L., a flood-tolerant, multiple-stemmed tree, which is restricted to the shoreline of BCI. Data on tree biometrics, epiphyte species, and epiphyte abundances were collected for more than 1200 trees. Results In total, we encountered almost 15,000 epiphytic individuals in sixty-eight species, corresponding to more than one third of the entire epiphyte flora of Barro Colorado Island. The component species differed strongly in abundance: the four most important species accounted for >75% of all individuals. In most cases, the same four species were also the first to colonize a tree (=phorophyte). Colonization patterns indicated no replacement of early colonizers by late arrivals. Species richness and epiphyte abundances showed a positive correlation with the size and the density of the host trees. All species showed a highly clumped distribution and the physiognomy of epiphyte communities of individual trees was dominated either by one or several of the four most common species or by a set of frequently co-occurring tank bromeliads. Other species were dominant only in exceptional cases. Most species were always rare. A distance effect on community composition was mostly confined to a local scale with an increased similarity in the species assemblage of stems of a tree v. neighbouring trees. Main conclusions The epiphytes on a single small phorophyte species may encompass a surprisingly large proportion of the local epiphyte flora. The observations that most tree crowns are inhabited by a single or only very few species, and that all epiphyte species show highly clumped distributions suggest a predominance of very local dispersal within a tree crown, which is only infrequently interrupted by successful long-distance dispersal between crowns.     

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.     

Linking woodland key habitat inventory and forest inventory data to prioritize districts needing conservation efforts

Woodland key habitat (WKH) inventories have been conducted in northern European countries, with the aim to create networks of minimally disturbed forest stands for protection. The goal of national forest inventory is to provide information relevant to forest management, such as on forest types, trees species composition, age structure and wood volume. The aim of this study was to link these two inventory databases to identify districts of Latvia most deficient in connectivity and habitat quality, in order to prioritize districts needing conservation effort. As an example, the area of deciduous forest with nemoral tree species (oak, ash, lime, maple and elm) and aspen was chosen. These forests provide habitat for a specific community of epiphytes. Using information in the WKH database, habitat quality in different districts of Latvia was estimated by the frequencies of occurrence of structural elements and selected indicator epiphyte species in nemoral tree species and aspen WKHs. Using digital data in the national forest inventory database, fragmentation metrics were determined for forests that, according to age and tree species composition, could potentially be nemoral tree and aspen WKHs. On a regional level, the lowest habitat quality in WKH occurred in districts that had the least fragmentation of potential WKH forest. In the less fragmented areas, the habitat quality of the existing WKH will likely increase in the future, and could be promoted by management to create structural elements typical of natural forests. The districts with the most fragmented nemoral and aspen forests, contained WKHs with the best habitat quality. A focus on protection should be given to these stands as they are the most likely to support source populations, and there is a need to improve spatial continuity of suitable tree substrate in these areas.