A forest habitat network for the Atlantic Oakwoods in Highland region,Scotland

Summary

The need to conserve biological diversity and ensure the future viability and integrity of Atlantic oakwoods in a fragmented landscape has led to strategies that facilitate a more holistic view of biodiversity conservation across extensive areas. The Scottish Forestry Strategy contains a major aspiration to develop forest habitat networks through the restoration and improvement of existing woodland and the expansion of new woodland. The Forest Research landscape ecology model BEETLE (Biological and Environmental Evaluation Tools for Landscape Ecology) uses a focal species approach to assess the functional connectivity of habitat within the wider landscape matrix. This model has been used to predict the current habitat network for Atlantic oakwood specialists in the Highland region of Scotland. The analysis outputs are presented with an approach to help practitioners and planners visualise the opportunities to target expansion, conversion or restoration of Atlantic oakwoods and their intrinsic biodiversity.     

Relative value of riparian woodlands in landscapes with different forest cover for medium-sized Iberian carnivores

This study analyses the role of riparian woodland in the conservation of five common carnivore species as compared with other non-riparian habitats according to woodland cover on the landscape scale (>60, 20–35 and <15% on 20 × 20-km surface area). I hypothesised that the importance of riparian woodlands in carnivore conservation on the regional scale would be greater in landscapes with low forest cover than in those with intermediate or high forest cover. To test this hypothesis, in each landscape type I sampled five riparian forests and 10–15 non-riparian habitats and recorded species richness and frequency of occurrence (number of sampled sites with species presence/total sampled sites) in both habitat types. The presence of (or use by) species at each particular sampling site was recorded using sign-surveys (search for scats and badger dens). The relative importance of riparian and non-riparian habitats in each landscape type was analysed by comparing species richness and frequency of occurrence for each species. Comparison of species richness and frequency of occurrence between landscape types indicate that riparian woodlands are important habitats in all landscape contexts. However, in accordance with the initial hypothesis, riparian woodlands are essential for carnivore conservation in the most deforested areas. Any agricultural or development policy should take into account the need to maintain and preserve riparian woodlands in landscapes, especially in intensively cultivated landscapes.     

森林所有制对景观格局和动物生境的影响研究进展

在欧美等地区的林业发达国家,多种森林所有制长期并存,至今,多种所有制影响下森林景观和生境格局的变化以及与动物保护之间的关系成为研究热点和重点.本文首先阐述了森林所有制对森林景观格局和动物生境的影响,以及林权分散和林权流转的加剧带来的生态后果,包括提供多样化的生境和导致原有大面积生境的破碎化;其次,总结了林业发达国家解决林权分散与生物保护之间矛盾的两个理论途径--改变所有制格局和相对统一各所有制的管理行为,分析现多采用后一种途径的原因,并指出目前森林所有制研究中模拟方法的局限;最后,论述我国的研究现状以及面临的机遇和挑战,提出近期发展建议.     

Lines in the land: a review of evidence for eastern Australia's major biogeographical barriers to closed forest taxa

The influence of climatic changes occurring since the late Miocene on Australia's eastern mesic ecosystems has received significant attention over the past 20 years. In particular, the impact of the dramatic shift from widespread rainforest habitat to a much drier landscape in which closed forest refugia were dissected by open woodland/savannah ecosystems has long been a focal point in Australian ecology and biogeography. Several specific regions along the eastern coast have been identified previously as potentially representing major biogeographical disjunctions for closed forest taxa. Initially, evidence stemmed from recognition of common zones where avian species/subspecies distributions and/or floral communities were consistently separated, but the body of work has since grown significantly with the rise of molecular phylogeographic tools and there is now a significant literature base that discusses the drivers, processes and effects of these hypothesised major biogeographical junctions (termed barriers). Here, we review the literature concerning eight major barriers argued to have influenced closed forest taxa; namely, the Laura Basin, Black Mountain Corridor, Burdekin Gap, Saint Lawrence Gap, Brisbane Valley Barrier, Hunter Valley Barrier, Southern Transition Zone and East Gippsland Barrier. We synthesise reported phylogeographical patterns and the inferred timing of influence with current climatic, vegetation and geological characteristics for each barrier to provide insights into regional evolution and seek to elicit common trends. All eight putative biogeographical barriers are characterised currently by lowland zones of drier, warmer, more open woodland and savannah habitat, with adjacent closed forest habitats isolated to upland cool, wet refugia. Molecular divergence estimates suggest two pulses of divergence, one in the early Miocene (~20–15 Mya) and a later one from the Pliocene–Pleistocene (~6–0.04 Mya). We conclude with a prospectus for future research on the eastern Australian closed forests and highlight critical issues for ongoing studies of biogeographical barriers worldwide.     

Using forest history and spatial patterns to identify potential high conservation value forests in Romania

Naturally dynamic forests have a high proportion of biotopes with old large trees, diverse vertical and horizontal structure at multiple scales, and much dead wood. As such, they provide habitat to species and ecosystem processes that forests managed for wood production cannot provide to the same degree. Whether termed old-growth, ancient, virgin, intact, primeval or continuity forests, a major challenge and need is to map such potential high conservation value forest for subsequent inclusion in functional habitat networks for biodiversity conservation in forest landscapes. Given that the delivery time of natural forest properties is much longer than of industry wood, we explore the usefulness of using historical maps to identify forests that have been continuously present for 220 years (potential old-growth) versus 140 years (potential aging forest) in a case study in the Romanian Carpathian Mountains (see Online Resource 1). While the total forest cover increased by 35 % over the past two centuries, the area of potential aging and potential old-growth forest declined by 56 and 34 %, respectively. Spatial modelling of edge effects and patch size for virtual species with different requirements indicated an even greater decrease in the area of functional habitat networks of old-growth and ageing forest. Our analyses show that compared to simple mapping of potential high conservation forests, the area of functional habitat patches is severely overestimated, and caution is needed when estimating the area of potential high conservation value forests that form functional habitat networks, i.e. a green infrastructure. In addition, the landscape and regional scale connectivity of patches needs to be considered. We argue that the use of historical maps combined with assessment of spatial patterns is an effective tool for identifying and analyzing potential high conservation value forests in a landscape context.     

A proposed strategy for maintaining mature forest habitat in Tasmania's wood production forests

Mature forests have structural habitat features that can take hundreds of years to develop, and large reserves alone are unlikely to ensure conservation of the species that rely on these features. This paper outlines a proposed new approach to managing mature forest features, the ‘mature habitat management approach’, in areas outside of reserves. The objective was to maintain a network of current and future mature forest habitat distributed across the landscape. The approach is designed to complement the existing reserve network and management actions and is tenure‐blind. Spatial information on the availability of mature forest habitat at the local (1‐km radius) and landscape (5‐km radius) scales is used for decisions on retention within a 1‐km radius of a harvest area, to reach the minimum target of 20% and 30% retention of mature forest at the local and landscape spatial scales, respectively. We believe this approach could contribute to meeting the conservation needs of many species that require mature forest features for refuge and breeding in Tasmania and elsewhere.     

Towards more effective integration of tropical forest restoration and conservation

Conservation and restoration interventions can be mutually reinforcing and are converging through an increased focus on social dimensions. This paper examines how to more effectively integrate the complementary goals of conservation and restoration of tropical forests. Forest conservation and restoration interventions are integral components of a broad approach to forest ecosystem and landscape management that aims to maintain and restore key ecological processes and enhance human well‐being, while minimizing biodiversity loss. The forest transition model provides a useful framework for understanding the relative importance of forest conservation and restoration interventions in different regions. Harmonizing conservation and restoration presents serious challenges for forest policy in tropical countries, particularly regarding the use and management of secondary forests, fallow vegetation, and forests degraded by logging and fire. Research to implement restoration more effectively in tropical regions can be stimulated by transforming questions that initially focused on conservation issues. Examination of papers published in Biotropica from 2000–2018 shows that most studies relevant to tropical forest conservation do not address forest restoration issues. Forest restoration studies, on the other hand, show a consistent association with conservation issues. There is much scope for further integration of conservation and restoration in research, practice, and policy. Securing a sustainable future for tropical forests requires developing and applying integrated approaches to landscape management that effectively combine knowledge and tools from multiple disciplines with practical experience and engagement of local stakeholders. Abstract in Portuguese is available with online material.     

Woodland restoration and forest structure affect nightjar abundance in the Ozark Highlands

Chuck-will's-widow (Antrostomus carolinensis) and eastern whip-poor-will (Antrostomus vociferus) are nightjars in eastern North America that have declined 69% and 67%, respectively, in abundance since 1966, resulting in conservation concerns for these species. We investigated relationships between nightjar abundance and landscape composition, forest structure, and application of tree thinning and prescribed fire because of regional interest in woodland restoration and nightjar conservation. We conducted nocturnal nightjar surveys at 385 points in southern Missouri, USA, in 2014 and 2015 and related counts to pine (Pinus spp.) and hardwood basal area, canopy closure, percent forest cover, and percent of area thinned or burned within 500 m of survey points. We modeled abundance of chuck-will's-widow and eastern whip-poor-will using time-removal models that included a detection process and an abundance process within a hierarchical Bayesian framework. We detected 534 eastern whip-poor-will and 186 chuck-will's-widow during surveys. Our data supported global models that included all 6 vegetation and management variables for both species. Chuck-will's-widow abundance was negatively related to hardwood basal area and peaked at intermediate values of percent area burned and percent forest cover. Eastern whip-poor-will abundance was negatively related to hardwood basal area and canopy cover, positively related to percent forest cover and percent of area burned, and peaked at low to moderate levels of percent of area thinned. Relationships to forest structure and management activities generally supported the conclusion that woodland restoration benefits nightjars and that chuck-will's-widow select landscapes with less forest cover than eastern whip-poor-will.     

道路对林地景观连接度的影响——以巩义市为例

基于景观连接度原理,借用景观连接度指数,在地理信息系统支持下,探讨了巩义市山区林地景观在不同距离阈值下连接度的变化,定量分析了道路对林地景观连接度的影响。结果显示,随着景观距离阈值的增大,无论是否有道路,林地景观整体可能连通性指数值都表现为逐渐增大;对林地景观连接度起"非常高"和"高"作用的林地斑块数量都比较少,但占林地总面积比例较大,面积大的林地斑块在提高景观连接度中起的作用较大;道路的分割使得林地斑块重要值降低,就单一斑块而言,随着景观距离阈值的增大,分割成的小斑块的重要值降低程度在逐渐减小。     

Dispersing male Parnassius smintheus butterflies are more strongly affected by forest matrix than are females

Dispersal is a central aspect of the ecology, evolution, and conservation of species. Predicting how species will respond to changing environmental conditions requires understanding factors that produce variation in dispersal. We explore one source of variation, differences between sexes within a spatial population network. Here, we compare the dispersal patterns of male and female Parnassius smintheus among 18 subpopulations over 8 years using the Virtual Migration Model. Estimated dispersal parameters differed between males and females, particularly with respect to movement through meadow and forest matrix habitat. The estimated dispersal distances of males through forest were much less than for females. Observations of female movement showed that, unlike males, females do not avoid forest nor does forest exert an edge effect. We explored whether further forest encroachment in this system would have different effects for males and females by fitting mean parameter estimates to the landscape configuration seen in 1993 and 2012. Despite differences in their dispersal due presumably to both habitat and physiological differences, males and females are predicted to respond in similar ways to reduced meadow area and increased forest isolation.     

Impacts of Restored Patch Density and Distance from Natural Forests on Colonization Success

Abstract The reduction and fragmentation of forest habitats is expected to have profound effects on plant species diversity as a consequence of the decreased area and increased isolation of the remnant patches. To stop the ongoing process of forest fragmentation, much attention has been given recently to the restoration of forest habitat. The present study investigates restoration possibilities of recently established patches with respect to their geographical isolation. Because seed dispersal events over 100 m are considered to be of long distance, a threshold value of 100 m between recent and old woodland was chosen to define isolation. Total species richness, individual patch species richness, frequency distributions in species occurrences, and patch occupancy patterns of individual species were significantly different among isolated and nonisolated stands. In the short term no high species richness is to be expected in isolated stands. Establishing new forests adjacent to existing woodland ensures higher survival probabilities of existing populations. In the long term, however, the importance of long‐distance seed dispersal should not be underestimated because most species showed occasional long‐distance seed dispersal. A clear distinction should be made between populations colonizing adjacent patches and patches isolated from old woodland. The colonization of isolated stands may have important effects on the dynamics and diversity of forest networks, and more attention should be directed toward the genetic traits and viability of founding populations in isolated stands.     

An overview of the ecology,management and conservation of Australia’s temperate woodlands

Summary Australia’s temperate woodlands are environments of cultural and ecological importance and significant repositories of Australia’s biodiversity. Despite this, they have been heavily cleared, much remaining vegetation is in poor condition and many species of plants and animals are threatened. Here, we provide a brief overview of key issues relating to the ecology, management and policy directions for temperate woodlands, by identifying and discussing ten themes. When addressing issues relating to the conservation and management of temperate woodlands, spatial scale is very important, as are the needs for a temporal perspective and a complementary understanding of pattern and process. The extent of landscape change in many woodland environments means that woodland patches, linear networks and paddock trees are critical elements, and that there can be pervasive effects from ‘problem’ native species such as the Noisy Miner (Manorina melanocephala). These consequences of landscape change highlight the challenge to undertake active management and restoration as well as effective monitoring and long‐term data collection. In developing approaches for conservation and management of temperate woodlands, it is essential to move our thinking beyond reserves to woodland conservation and management on private land, and recognise the criticality of cross‐disciplinary linkages. We conclude by identifying some emerging issues in woodland conservation and management. These include the need to further develop non‐traditional approaches to conservation particularly off‐reserve management; the value of documenting approaches and programmes that demonstrably lead to effective change; new lessons that can be learned from intact examples of temperate woodlands; and the need to recognise how climate change and human population growth will interact with conservation and management of temperate woodlands in future decades.     

Humid tropical rain forest has expanded into eucalypt forest and savanna over the last 50 years

Tropical rain forest expansion and savanna woody vegetation thickening appear to be a global trend, but there remains uncertainty about whether there is a common set of global drivers. Using geographic information techniques, we analyzed aerial photography of five areas in the humid tropics of northeastern Queensland, Australia, taken in the 1950s and 2008, to determine if changes in rain forest extent match those reported for the Australian monsoon tropics using similar techniques. Mapping of the 1950s aerial photography showed that of the combined study area (64,430 ha), 63% was classified as eucalypt forests/woodland and 37% as rain forest. Our mapping revealed that although most boundaries remained stable, there was a net increase of 732 ha of the original rain forest area over the study period, and negligible conversion of rain forest to eucalypt forest/woodland. Statistical modeling, controlling for spatial autocorrelation, indicated distance from preexisting rain forest as the strongest determinant of rain forest expansion. Margin extension had a mean rate across the five sites of 0.6 m per decade. Expansion was greater in tall open forest types but also occurred in shorter, more flammable woodland vegetation types. No correlations were detected with other local variables (aspect, elevation, geology, topography, drainage). Using a geographically weighted mean rate of rain forest margin extension across the whole region, we predict that over 25% of tall open forest (a forest type of high conservation significance) would still remain after 2000 years of rain forest expansion. This slow replacement is due to the convoluted nature of the rain forest boundary and the irregular shape of the tall open forest patches. Our analyses point to the increased concentration of atmospheric CO(2) as the most likely global driver of indiscriminate rain forest expansion occurring in northeastern Australia, by increasing tree growth and thereby overriding the effects of fire disturbance.     

Quantitative changes of forest landscapes over the last century across Italy

A key topic in landscape ecology and vegetation science is the quantitative analysis of changes in forest cover over time, through the use of geomatics monitoring tools. Ecologists and landscape researchers are pointing out that a full understanding of ecosystems and landscapes should be based on the analysis of their functioning over long time series. Under this perspective, a long-term historical reconstruction of forest cover is essential. This study has aimed at examining the long-term dynamics of forest landscapes in Italy, over the last century, using recent remote-sensing based map (2012) and an accurate historical map (1936). A forest-non forest approach has been followed by the computation of a variety of landscape metrics using two analysis tools, with the final objective of quantifying changes in forest cover patterns and in the composition of specific landscape elements. Results show that forest landscape structure has significantly changed across Italy, resulting in a general trend of decreasing fragmentation and patchiness, mainly through enlargement of existing forest patches, which have also assumed a more geometrically regular shape. In relative terms, the greatest expansion of forest areas has occurred mainly in lowland districts characterised by the highest level of human pressure in the country.     

Using historical woodland creation to construct a long‐term,large‐scale natural experiment: the WrEN project

Natural experiments have been proposed as a way of complementing manipulative experiments to improve ecological understanding and guide management. There is a pressing need for evidence from such studies to inform a shift to landscape‐scale conservation, including the design of ecological networks. Although this shift has been widely embraced by conservation communities worldwide, the empirical evidence is limited and equivocal, and may be limiting effective conservation. We present principles for well‐designed natural experiments to inform landscape‐scale conservation and outline how they are being applied in the WrEN project, which is studying the effects of 160 years of woodland creation on biodiversity in UK landscapes. We describe the study areas and outline the systematic process used to select suitable historical woodland creation sites based on key site‐ and landscape‐scale variables – including size, age, and proximity to other woodland. We present the results of an analysis to explore variation in these variables across sites to test their suitability as a basis for a natural experiment. Our results confirm that this landscape satisfies the principles we have identified and provides an ideal study system for a long‐term, large‐scale natural experiment to explore how woodland biodiversity is affected by different site and landscape attributes. The WrEN sites are now being surveyed for a wide selection of species that are likely to respond differently to site‐ and landscape‐scale attributes and at different spatial and temporal scales. The results from WrEN will help develop detailed recommendations to guide landscape‐scale conservation, including the design of ecological networks. We also believe that the approach presented demonstrates the wider utility of well‐designed natural experiments to improve our understanding of ecological systems and inform policy and practice.     

Higher climate warming sensitivity of Siberian larch in small than large forest islands in the fragmented Mongolian forest steppe

Forest fragmentation has been found to affect biodiversity and ecosystem functioning in multiple ways. We asked whether forest size and isolation in fragmented woodlands influences the climate warming sensitivity of tree growth in the southern boreal forest of the Mongolian Larix sibirica forest steppe, a naturally fragmented woodland embedded in grassland, which is highly affected by warming, drought, and increasing anthropogenic forest destruction in recent time. We examined the influence of stand size and stand isolation on the growth performance of larch in forests of four different size classes located in a woodland‐dominated forest‐steppe area and small forest patches in a grassland‐dominated area. We found increasing climate sensitivity and decreasing first‐order autocorrelation of annual stemwood increment with decreasing stand size. Stemwood increment increased with previous year's June and August precipitation in the three smallest forest size classes, but not in the largest forests. In the grassland‐dominated area, the tree growth dependence on summer rainfall was highest. Missing ring frequency has strongly increased since the 1970s in small, but not in large forests. In the grassland‐dominated area, the increase was much greater than in the forest‐dominated landscape. Forest regeneration decreased with decreasing stand size and was scarce or absent in the smallest forests. Our results suggest that the larch trees in small and isolated forest patches are far more susceptible to climate warming than in large continuous forests pointing to a grim future for the forests in this strongly warming region of the boreal forest that is also under high land use pressure.     

Impact of landscape fragmentation on a specialised woodland bat,Rhinolophus hipposideros

For habitat specialists, fragmentation has major consequences as it means less suitable habitat for the species to live in. In a fragmented landscape, we would expect larger, but spatially more clustered, foraging ranges. We studied the impact of landscape fragmentation on the foraging range and habitat exploitation of a specialised forest bat by radiotracking 16 female lesser horseshoe bats Rhinolophus hipposideros in a landscape with connected woodland structures and in a highly fragmented landscape in Carinthia, Austria.Contrary to our expectations, spatial foraging behaviour was not influenced by fragmentation. No differences in the behaviour of the bats between the sites were evident for the foraging ranges (minimum convex polygon, MCP), the core foraging areas (50% kernel), nor the mean or the maximum distances from the roost. However, in the highly fragmented landscape, the foraging activity of individuals was spatially more clustered and the overall MCP of all bats of a colony was greater compared to the less fragmented landscape.Woodland was the most important foraging habitat for the lesser horseshoe bats at both study sites. Habitat selection at the individual MCPs was evident only at the site with low fragmentation. However, in the core foraging areas, woodland was significantly selected over all other habitat types at both study sites.We conclude that (1) conservation measures for colonies of lesser horseshoe bats should be undertaken within 2.5 km of the nursery roost, (2) woodland is the key foraging habitat particularly in the vicinity of the roost, and (3) any loss of woodland near the colonial roosts are likely to negatively influence the colony, since these bats do not seem to be able to adapt their spatial foraging behaviour in a degraded landscape. The inflexible spatial behaviour of this specialised bat highlights the need to compensate for any habitat loss within the foraging range of a bat colony.     

Natural forest regeneration and ecological restoration in human‐modified tropical landscapes

In human‐modified tropical landscapes (HMLs) the conservation of biodiversity, functions and services of forest ecosystems depends on persistence of old growth forest remnants, forest regeneration in abandoned agricultural fields, and restoration of degraded lands. Understanding the impacts of agricultural land uses (ALUs) on forest regeneration is critical for biodiversity conservation in HMLs. Here, we develop a conceptual framework that considers the availability of propagules and the environment prevailing after field abandonment as two major determinants of forest regeneration in HMLs. The framework proposes that regeneration potential decreases with size, duration and severity of agricultural disturbance, reducing propagule availability and creating ill‐suited environmental conditions for regeneration. We used studies from Southern Mexico to assess this framework. First, we identify regeneration bottlenecks that trees face during transit from seed to follow‐up life stages, using demographic analysis of dominant pioneer species in recently abandoned fields. Then, we explore effects of ALUs on forest regeneration at the field and landscape scales, addressing major legacies. Finally, we integrate agricultural disturbance with landscape composition to predict attributes of successful second growth forests in HMLs, and provide indicators useful to select tree native species for active restoration. An indicator of disturbance inflicted by ALUs, based on farmers’ information, predicted better regeneration potential than measurements of soil and microclimate conditions at time of abandonment. Cover of cattle pastures in the landscape was a stronger indicator of forest regenerating attributes than cover of old growth forest remnants. To conclude, we offer recommendations to promote forest regeneration and biodiversity conservation in HMLs.     

Spatial variation and biogeography of sand forest avian assemblages in South Africa

Aim To examine biogeographical affiliations, habitat‐associated heterogeneity and endemism of avian assemblages in sand forest patches and the savanna‐like mixed woodland matrix. Location Two reserves in the Maputaland Centre of Endemism (MC) on the southern Mozambique Coastal Plain of northern KwaZulu‐Natal, South Africa. Methods Replicated surveys were undertaken in each of the two habitat types in each reserve, providing species abundance data over a full year. Vegetation structure at each of the survey sites was also quantified. Differences between the bird assemblages and the extent to which vegetation structure explained these differences were assessed using multi‐variate techniques. Biogeographical comparisons were based on species presence/absence data and clustering techniques. Results Bird assemblages differed significantly between habitats both within a given reserve and between reserves, and also between reserves for a given habitat. Differences in vegetation structure contributed substantially to differences between the avian assemblages. Of the four species endemic to the MC, three (Neergaard’s sunbird, Rudd’s apalis, and Woodward’s batis) were consistently present in sand forest. The fourth (pink‐throated twinspot) preferred mixed woodland. None of these endemic species was classed as rare. In the biogeographical analysis, both the sand forest and the mixed woodland bird assemblages were most similar to bird assemblages found in the forest biome or the Afromontane forest biome, depending on the biome classification used. Main conclusions The close affinities of sand forest and mixed woodland assemblages to those of the forest biome are most likely due to similarities in vegetation structure of these forests. Bird assemblages differ between the sand forest and mixed woodland habitats both within a given reserve and between reserves, and also between reserves for a given habitat. These differences extend to species endemic to the MC. Thus, conservation of sand forest habitat in a variety of areas is necessary to ensure the long‐term persistence of the biota.     

Structures determining bryophyte species richness in a managed forest landscape in boreo-nemoral Europe

Forest patches with high biological value are protected as woodland key habitats (WKH), which are identified by the presence of forest structures and indicator species. However, management for conservation needs to consider also managed forests as habitats for species. In this respect, there is a need to set quantitative targets for species and structures at different landscape scales. Due to non-intensive methods of forest management used prior to 1940 in Latvia, it might be expected that large areas of forest have developed structures that can support many species characteristic of natural forests. The aim of the study was to create a model that best described the richness of bryophyte species that are characteristic of natural forests, using forest structures as explanatory factors. The structures and bryophyte communities on living trees and coarse woody debris (CWD) were described in plots along transects blindly placed in areas dominated by State forests under commercial management. Explanatory variables related to tree species composition and tree size explained 54% of the variation in WKH indicator species richness on living trees. The best explanatory factors were maximum diameter of deciduous tree species and CWD. Low richness of total bryophyte and indicator species was found on dead wood, and the amount of variation in bryophyte species richness on CWD explained by explanatory variables was low. The study indicates the importance of deciduous tree substrate in managed forests in maintaining the spatial continuity of epiphytic species diversity. However, the forests in the managed forest landscape did not support high diversity of epixylic species, even in the WKHs, due to low diversity of suitable dead wood substrate.