Colonization–extinction dynamics of epixylic lichens along a decay gradient in a dynamic landscape

Metapopulation models are often used for understanding and predicting species dynamics in fragmented landscapes. Several models have been proposed depending on e.g. the relative importance of patch dynamics on the metapopulation dynamics. Dead wood is a dynamic substrate patch, and species that are confined to such patches have experienced a high degree of habitat loss in managed forests. Little is, however, known about how the population dynamics of epixylic species are affected by the fast dynamics of their substrate patches. We quantified the effect of local patch conditions and metapopulation processes on colonizations and extinctions of epixylic lichen species in a managed boreal forest landscape. This was done by twice surveying seven lichen metapopulations on 293 stumps in 30 stands of ages covering the duration of the dynamic patches (stumps). We also investigated the relative importance of local stochastic extinctions from stumps that remained available, and deterministic extinctions due to stump surface disappearance. We found importance of a decay gradient, surrounding metapopulation size, and local population sizes, in driving the colonization–extinction dynamics of epixylic lichens. The species were sorted along the stump decay gradient. Increasing surrounding metapopulation size was associated with increased colonization rates, and increasing local population size decreased lichen extinction rates. Finally, both local stochastic extinctions and deterministic extinctions due to patch disappearance occur, confirming that the long‐term persistence of epixylic lichens depends on colonization rates that compensate for stochastic population extinctions as well as deterministic extinctions.     

Epiphytic macrolichens in managed and natural forest landscapes: a comparison at two spatial scales

To maintain biodiversity in managed forests we must understand how forestry affects various organisms across a wide range of spatial and temporal scales. We compared landscape structure, forest structure, and species richness and abundance of epiphytic macrolichens in three pairs of natural and managed boreal forest landscapes. Study landscapes (2500 ha) were located within and adjacent to three of the largest forest reserves in Sweden (Reivo, Muddus, Jelka). The structural heterogeneity within landscapes was higher in managed forests whereas within-stand structural heterogeneity was higher in natural landscapes. Species richness of macrolichens at the stand level (sample plot) was 23% higher in natural forests but there was no difference at the landscape level. Most (86%) of the common species were more frequent in natural landscapes. Lichen abundance (estimated by lichen litter) was two times higher in natural than in managed landscapes, 5.6 and 2.7 kg ha^-1 forest (pooled data), respectively. Both species richness and abundance were negatively related to cutting level (number and basal area of cut stumps) and positively related to stand variables (stand age, stem density and basal area). Lichen-rich forest stands were more numerous but covered a smaller area and were more isolated in managed landscapes. This may in turn have important consequences for dispersal of lichen propagules to second-growth forests. In conclusion, the results suggest that effects of forestry on epiphyte diversity and abundance are strongly related to the spatial scale (stand or landscape). To enhance biodiversity in managed forests we must increase structural heterogeneity at the whole range of spatial and temporal scales.     

Biodiversity and ecological forest-cover domains in boreal landscapes

Studies of spatial forest patterns have traditionally focused on habitat remnants within a landscape, but few have explicitly accounted for natural habitat patterns inherent to those landscapes. At broader scales, all cover types are to some degree subdivided. In boreal forest landscapes, forests, peat lands and water bodies form a variety of different spatial structures depending on their proportions and arrangement. To assess the spatial arrangement of forest patches relevant to organisms and ecological processes, we systematically sampled and analysed 57 boreal forest landscapes (10,000 ha in size) in central Finland. Our results show that even though forest is spatially very subdivided into discrete patches in boreal landscapes it becomes generally well-connected if narrow non-forested gaps are ignored. The proportion of forest cover varied from 17.8 to 75.3% and the number of discrete forest patches from 37 to 213. The average percolation threshold for forest cover was 46.8%. If ≤100 m wide non-forested gaps were ignored and forest patches were joined, the percolation threshold dropped to 33.3%. There were on average 13 discrete clusters of forest patches if forest patches within 200 m distance were combined. These results suggest that if a boreal forest species is able to cross even relatively narrow non-forested gaps, it is likely to perceive these boreal forest landscapes as continuous. Even though our present analysis was based on static forest cover patterns, it is important to consider landscape pattern domains when assessing habitat fragmentation and its consequences to populations of organisms.     

Naturally Saline Boreal Communities as Models for Reclamation of Saline Oil Sand Tailings

Reclaimed landscapes after oil sands mining have saline soils; yet, they are required to have similar biodiversity and productivity as the predisturbance nonsaline landscape. Given that many species in the boreal forest are not tolerant of salinity, we studied the effects of soil salinity on plant communities in natural saline landscapes to understand potential plant responses during the reclamation process. Vegetation–soil relationships were measured along transects from flooded wetlands to upland forest vegetation in strongly saline, slightly saline, nonsaline, and reclaimed boreal landscapes. In strongly saline landscapes, surface soil salinity was high (>10 dS/m) in flooded, wet‐meadow, and dry‐meadow vegetation zones as compared to slightly saline (<5 dS/m) and nonsaline (<2 dS/m) landscapes. Plant communities in these vegetation zones were quite different from nonsaline boreal landscapes and were dominated by halophytes common to saline habitats of the Great Plains. In the shrub and forest vegetation zones, surface soil salinity was similar between saline and nonsaline landscapes, resulting in similar plant communities. In strongly saline landscapes, soils remained saline at depth through the shrub and forest vegetation zones (>10 dS/m), suggesting that forest vegetation can establish over saline soils as long as the salts are below the rooting zone. The reclaimed landscape was intermediate between slightly saline and nonsaline landscapes in terms of soil salinity but more similar to nonsaline habitats with respect to species composition. Results from this study suggest it may be unrealistic to expect that plant communities similar to those found on the predisturbance landscape can be established on all reclaimed landscapes after oil sands mining.     

Landscape-variability of the carbon balance across managed boreal forests

Boreal forests are important global carbon (C) sinks and, therefore, considered as a key element in climate change mitigation policies. However, their actual C sink strength is uncertain and under debate, particularly for the actively managed forests in the boreal regions of Fennoscandia. In this study, we use an extensive set of biometric- and chamber-based C flux data collected in 50 forest stands (ranging from 5 to 211 years) over 3 years (2016–2018) with the aim to explore the variations of the annual net ecosystem production (NEP; i.e., the ecosystem C balance) across a 68 km² managed boreal forest landscape in northern Sweden. Our results demonstrate that net primary production rather than heterotrophic respiration regulated the spatio-temporal variations of NEP across the heterogeneous mosaic of the managed boreal forest landscape. We further find divergent successional patterns of NEP in our managed forests relative to naturally regenerating boreal forests, including (i) a fast recovery of the C sink function within the first decade after harvest due to the rapid establishment of a productive understory layer and (ii) a sustained C sink in old stands (131–211 years). We estimate that the rotation period for optimum C sequestration extends to 138 years, which over multiple rotations results in a long-term C sequestration rate of 86.5 t C ha⁻¹ per rotation. Our study highlights the potential of forest management to maximize C sequestration of boreal forest landscapes and associate climate change mitigation effects by developing strategies that optimize tree biomass production rather than heterotrophic soil C emissions.     

Saproxylic beetle diversity in a managed boreal forest: importance of stand characteristics and forestry conservation measures

Saproxylic beetles constitute a significant proportion of boreal forest biodiversity. However, the long history of timber production in Fennoscandia has significantly reduced the availability of dead wood and is considered a threat to the conservation of saproxylic beetle assemblages. Therefore, since the mid‐1990s dead wood retention in harvested stands has formed an integral part of silvicultural practices. However, the contribution of this biodiversity‐orientated management approach to conserving saproxylic beetle assemblages in boreal forest landscapes that include production forestry remains largely untested. We examined differences in resident saproxylic beetle assemblages among stands under different management in a boreal forest landscape in Central Sweden, and in particular stands managed according to new conservation‐orientated practices. We also investigated the relationship between beetle diversity and forest stand characteristics. Bark of coarse woody debris (CWD) was sieved for beetles in old managed stands, unmanaged nature reserves, and set‐aside areas, and clear‐cut stands harvested according to certification guidelines [new forestry (NF) clear‐cuts]. All stand types contributed significantly to the total diversity of beetles found. While stand size, position, and distance to nearest reserve were unimportant, both the quality and the quantity of CWD in stands contributed significantly to explaining beetle abundance and species richness. This extends the previous findings for red‐listed invertebrates, and shows that heterogeneous substrate quality and a range of management practices are necessary to maintain saproxylic beetle diversity in boreal forest landscapes that include production forestry. The unique abiotic conditions in combination with the abundant and varied CWD associated with NF clear‐cuts form an important component of forest stand heterogeneity for saproxylic beetles. It is thus essential that sufficient, diverse, CWD is retained in managed boreal landscapes to ensure the conservation of boreal saproxylic beetle assemblages.     

Patterns and Dynamics of Dissolved Organic Carbon (DOC) in Boreal Streams: The Role of Processes,Connectivity, and Scaling

We bring together three decades of research from a boreal catchment to facilitate an improved mechanistic understanding of surface water dissolved organic carbon (DOC) regulation across multiple scales. The Krycklan Catchment Study encompasses 15 monitored nested research catchments, ranging from 3 to 6900 ha in size, as well as a set of monitored transects of forested and wetland soils. We show that in small homogenous catchments, hydrological functioning provides a first order control on the temporal variability of stream water DOC. In larger, more heterogeneous catchments, stream water DOC dynamics are regulated by the combined effect of hydrological mechanisms and the proportion of major landscape elements, such as wetland and forested areas. As a consequence, streams with heterogeneous catchments undergo a temporal switch in the DOC source. In a typical boreal catchment covered by 10-20% wetlands, DOC originates predominantly from wetland sources during low flow conditions. During high flow, the major source of DOC is from forested areas of the catchment. We demonstrate that by connecting knowledge about DOC sources in the landscape with detailed hydrological process understanding, an improved representation of stream water DOC regulation can be provided. The purpose of this study is to serve as a framework for appreciating the role of regulating mechanisms, connectivity and scaling for understanding the pattern and dynamics of surface water DOC across complex landscapes. The results from this study suggest that the sensitivity of stream water DOC in the boreal landscape ultimately depends on changes within individual landscape elements, the proportion and connectivity of these affected landscape elements, and how these changes are propagated downstream.     

The effect of forest management operations on population performance of Vaccinium myrtillus on a landscape-scale

We have studied the variation in average performance of Vaccinium myrtillus L. among local populations within an intensively managed boreal forest landscape in southeast Norway. Our aim was to identify whether forest disturbance history was an important factor controlling the variation in average plant performance. We used three measures of forest maturity (maturity class, tree biomass and number of trees) as variables reflecting disturbance history. These variables were used in our analysis together with measured environmental conditions (soil moisture, pH, nitrogen, and elevation) a priori known to affect plant performance. To quantify plant performance, we used multiple measures of the local V. myrtillus populations, including percentage cover, average ramet age, height, diameter, biomass, growth, and sexual reproduction (number of fruits and number of mature seeds per fruit). Most of the explained variation in our performance variables could be related to our environmental variables, but disturbance history was also an important determinant of the performance of V. myrtillus. All performance variables were higher in younger, less mature stands, except ramet age, which was highest in mature stands. The increase in performance after clear cutting indicates that this period is of high importance for the population dynamics and persistence of V. myrtillus in intensively managed boreal forest landscapes.     

Landscape-Scale Disturbances Modified Bird Community Dynamics in Successional Forest Environment

Ecosystem-based forest management strives to develop silvicultural practices that best emulate natural disturbances such as wildfire to conserve biodiversity representative of natural forest ecosystems. Yet, current logging practices alter forest structure and reduce the proportion of old-growth forest and, consequently, can exert long-term effects on the dynamics of forest biota. The stand- and landscape-scale factors driving bird community dynamics in post-disturbance environment remain poorly understood. In this study, we examined bird community dynamics along successional gradients in boreal ecosystems originating from fire and logging in landscapes dominated by old-growth forest. We tested if bird species richness and community compositions in clear-cutting stands became comparable to those in natural stands after 70 years, and identified the relative contributions of stand- and landscape-scale forest attributes in bird community dynamics. Based on records of bird occurrences at 185 field sites in natural and clearcutting stands, we demonstrate that (1) both forest structures and bird communities underwent evident changes along successional gradients in post-clearcutting environment; (2) bird species richness and community composition in 60- to 70-years-old clearcutting stands still differed from those in 50- to 79-years-old natural stands, in spite of the fact that most forest attributes of clearcutting stands became comparable to those of natural stands after 40 years; and (3) landscape disturbances contributed more than stand characteristics in explaining the lack of convergence of mature forest species, residents, and short-distance migrants in post-clearcutting environment. Our study points out that more regards should be paid to improve the landscape configuration of the managed forests, and implies that old-growth forest retention within logged areas, combined with selection cutting and prolonged logging rotations, can better emulate fire and alleviate forest harvesting effects on bird community assemblages typical of natural boreal ecosystem.     

Effects of the landscape on boreal toad gene flow: does the pattern–process relationship hold true across distinct landscapes at the northern range margin?

Understanding the impact of natural and anthropogenic landscape features on population connectivity is a major goal in evolutionary ecology and conservation. Discovery of dispersal barriers is important for predicting population responses to landscape and environmental changes, particularly for populations at geographic range margins. We used a landscape genetics approach to quantify the effects of landscape features on gene flow and connectivity of boreal toad (Bufo boreas) populations from two distinct landscapes in south-east Alaska (Admiralty Island, ANM, and the Chilkat River Valley, CRV). We used two common methodologies for calculating resistance distances in landscape genetics studies (resistance based on least-cost paths and circuit theory). We found a strong effect of saltwater on genetic distance of CRV populations, but no landscape effects were found for the ANM populations. Our discordant results show the importance of examining multiple landscapes that differ in the variability of their features, to maximize detectability of underlying processes and allow results to be broadly applicable across regions. Saltwater serves as a physiological barrier to boreal toad gene flow and affects populations on a small geographic scale, yet there appear to be few other barriers to toad dispersal in this intact northern region.     

An improved neutral landscape model for recreating real landscapes and generating landscape series for spatial ecological simulations

Many studies have assessed the effect of landscape patterns on spatial ecological processes by simulating these processes in computer‐generated landscapes with varying composition and configuration. To generate such landscapes, various neutral landscape models have been developed. However, the limited set of landscape‐level pattern variables included in these models is often inadequate to generate landscapes that reflect real landscapes. In order to achieve more flexibility and variability in the generated landscapes patterns, a more complete set of class‐ and patch‐level pattern variables should be implemented in these models. These enhancements have been implemented in Landscape Generator (LG), which is a software that uses optimization algorithms to generate landscapes that match user‐defined target values. Developed for participatory spatial planning at small scale, we enhanced the usability of LG and demonstrated how it can be used for larger scale ecological studies. First, we used LG to recreate landscape patterns from a real landscape (i.e., a mountainous region in Switzerland). Second, we generated landscape series with incrementally changing pattern variables, which could be used in ecological simulation studies. We found that LG was able to recreate landscape patterns that approximate those of real landscapes. Furthermore, we successfully generated landscape series that would not have been possible with traditional neutral landscape models. LG is a promising novel approach for generating neutral landscapes and enables testing of new hypotheses regarding the influence of landscape patterns on ecological processes. LG is freely available online.     

Metapopulation dynamics of a beetle species confined to burned forest sites in a managed forest region

Despite increasing awareness of the theoretical importance of habitat dynamics on metapopulations, only a few empirical studies have been conducted. We aimed to increase our understanding of how patch size, dynamics and connectivity affect colonization–extinction dynamics and the occurrence patterns of a beetle (Stephanopachys linearis), which breeds only in burned trees, existing as dynamic habitat patches that have become rare in managed forest landscapes. We assessed species’ presence/absence twice in all known habitat patches (i.e. > 1 ha sites where forest fires had occurred during the previous 2–15 yr) in a 200 × 150 km region of central Sweden, dominated by managed boreal forest. Evaluated over six years, the colonization rate was 47% and the local extinction risk was 65%. Probability of colonization increased with patch size (number of suitable trees in a site) and connectivity to occupied patches within 30 km, and decreased with increasing time since fire. Local extinction risk decreased with habitat patch size but increased, unexpectedly, with connectivity. Occurrence increased with patch size and decreased with increasing time since fire. At a regional scale, S. linearis tracks the fire dynamics by colonising sites with burned trees and by becoming extinct at rates which make the species rare at sites where burnt trees are more than eight years old. In managed boreal forest landscapes, a large proportion of sites may be created by prescribed burning (in our study area: 82%), and consequently human decisions strongly affect the future amount of habitat for fire‐dependent species and its spatial distribution. Stephanopachys linearis uses burned sites more often if more trees are retained and, to some extent, if sites are concentrated in those parts of a region that already support high population densities of the species.     

城市生态景观研究的基础理论框架与技术构架

构建人与自然和谐的生态景观是当今城市发展的重要目标之一,统一的内涵和完善的指标体系是城市生态景观规划和设计的迫切需求。通过概括城市生态景观的基本内涵和主要特征,认为其核心价值包括可持续性、生态经济效益及和谐性。据此,提出了8条指导原则,包括景观自然度、物种安全性和多样性、雨水利用效率、污染物排放和处理、绿色能源和材料利用、局地小气候调节、景观视觉享受以及精神内涵和文化品位,并根据文献资料和行业导则,整理并归纳了有关的定量评价指标和适用范围。最后总结了城市生态景观的技术构架,以及在资源保护和节约、功能与形式的有机结合、系统生态价值最大化等方面亟待解决的关键技术。为城市生态景观研究提供基础理论框架,也可以为城市生态景观的具体实践提供参考。     

Could increased boreal forest ecosystem productivity offset carbon losses from increased disturbances?

To understand how boreal forest carbon (C) dynamics might respond to anticipated climatic changes, we must consider two important processes. First, projected climatic changes are expected to increase the frequency of fire and other natural disturbances that would change the forest age-class structure and reduce forest C stocks at the landscape level. Second, global change may result in increased net primary production (NPP). Could higher NPP offset anticipated C losses resulting from increased disturbances? We used the Carbon Budget Model of the Canadian Forest Sector to simulate rate changes in disturbance, growth and decomposition on a hypothetical boreal forest landscape and to explore the impacts of these changes on landscape-level forest C budgets. We found that significant increases in net ecosystem production (NEP) would be required to balance C losses from increased natural disturbance rates. Moreover, increases in NEP would have to be sustained over several decades and be widespread across the landscape. Increased NEP can only be realized when NPP is enhanced relative to heterotrophic respiration. This study indicates that boreal forest C stocks may decline as a result of climate change because it would be difficult for enhanced growth to offset C losses resulting from anticipated increases in disturbances.     

Explaining Geographic Gradients in Winter Selection of Landscapes by Boreal Caribou with Implications under Global Changes in Eastern Canada

Many animal species exhibit broad-scale latitudinal or longitudinal gradients in their response to biotic and abiotic components of their habitat. Although knowing the underlying mechanism of these patterns can be critical to the development of sound measures for the preservation or recovery of endangered species, few studies have yet identified which processes drive the existence of geographical gradients in habitat selection. Using extensive spatial data of broad latitudinal and longitudinal extent, we tested three hypotheses that could explain the presence of geographical gradients in landscape selection of the endangered boreal woodland caribou (Rangifer tarandus caribou) during winter in Eastern Canadian boreal forests: 1) climate-driven selection, which postulates that geographic gradients are surrogates for climatic gradients; 2) road-driven selection, which proposes that boreal caribou adjust their selection for certain habitat classes as a function of proximity to roads; and 3) an additive effect of both roads and climate. Our data strongly supported road-driven selection over climate influences. Thus, direct human alteration of landscapes drives boreal caribou distribution and should likely remain so until the climate changes sufficiently from present conditions. Boreal caribou avoided logged areas two-fold more strongly than burnt areas. Limiting the spread of road networks and accounting for the uneven impact of logging compared to wildfire should therefore be integral parts of any habitat management plan and conservation measures within the range of the endangered boreal caribou. The use of hierarchical spatial models allowed us to explore the distribution of spatially-structured errors in our models, which in turn provided valuable insights for generating alternative hypotheses about processes responsible for boreal caribou distribution.     

Importance of high quality early-successional habitats in managed forest landscapes to rare beetle species

Species adapted to early-successional forest habitats are in managed landscapes largely confined to clearcuts. To improve habitat quality on clearcuts, green tree and dead wood retention is widely applied in forestry; however, its effects on rare early-successional species have rarely been shown. We repeatedly surveyed two red-listed beetle species (Upis ceramboides and Platysoma minus) on clearcuts in a managed boreal forest landscape. We found that U. ceramboides decreased its occupancy over time while P. minus increased, indicating that red-listed species vary in their ability to successfully utilise managed habitats. We found no effect of connectivity on probability of occurrence, colonisation or extinction per clearcut. Trees retained alive improved habitat quality of clearcuts, since both species were more frequent in dead wood of such trees, in comparison to logging residues. We suggest that retention can be improved by protecting and creating dead wood as intact trees during harvesting. Rare specialist species require habitat of high quality, and consequently it is impossible to meet the requirements of these species on every clearcut. To preserve all early-successional species at a regional scale, we recommend focusing retention of green trees and dead wood to one or a few trees species on each clearcut and in each landscape.     

Landscape issues in plant ecology

In the last decade, we have seen the emergence and consolidation of a conceptual framework that recognizes the landscape as an ecological unit of interest. Plant ecologists have long emphasized landscape-scale issues, but there has been no recent attempt to define how landscape concepts are now integrated in vegetation studies. To help define common research paradigms in both landscape and plant ecology, we discuss issues related to three main landscape concepts in vegetation researches, reviewing theoretical influences and emphasizing recent developments. We first focus on environmental relationships, documenting how vegetation patterns emerge from the influence of local abiotic conditions. The landscape is the physical environment. Disturbances are then considered, with a particular attention to human-driven processes that often overrule natural dynamics. The landscape is a dynamic space. As environmental and historical processes generate heterogeneous patterns, we finally move on to stress current evidence relating spatial structure and vegetation dynamics. This relates to the concept of a landscape as a patch-corridor-matrix mosaic. Future challenges involve: 1) the capacity to evaluate the relative importance of multiple controlling processes at broad spatial scale; 2) better assessment of the real importance of the spatial configuration of landscape elements for plant species and finally; 3) the integration of natural and cultural processes and the recognition of their interdependence in relation to vegetation management issues in human landscapes.     

The Net Landscape Carbon Balance—Integrating terrestrial and aquatic carbon fluxes in a managed boreal forest landscape in Sweden

The boreal biome exchanges large amounts of carbon (C) and greenhouse gases (GHGs) with the atmosphere and thus significantly affects the global climate. A managed boreal landscape consists of various sinks and sources of carbon dioxide (CO₂), methane (CH₄), and dissolved organic and inorganic carbon (DOC and DIC) across forests, mires, lakes, and streams. Due to the spatial heterogeneity, large uncertainties exist regarding the net landscape carbon balance (NLCB). In this study, we compiled terrestrial and aquatic fluxes of CO₂, CH₄, DOC, DIC, and harvested C obtained from tall‐tower eddy covariance measurements, stream monitoring, and remote sensing of biomass stocks for an entire boreal catchment (~68 km²) in Sweden to estimate the NLCB across the land–water–atmosphere continuum. Our results showed that this managed boreal forest landscape was a net C sink (NLCB = 39 g C m^?2 year^?1) with the landscape–atmosphere CO₂ exchange being the dominant component, followed by the C export via harvest and streams. Accounting for the global warming potential of CH₄, the landscape was a GHG sink of 237 g CO₂‐eq m^?2 year^?1, thus providing a climate‐cooling effect. The CH₄ flux contribution to the annual GHG budget increased from 0.6% during spring to 3.2% during winter. The aquatic C loss was most significant during spring contributing 8% to the annual NLCB. We further found that abiotic controls (e.g., air temperature and incoming radiation) regulated the temporal variability of the NLCB whereas land cover types (e.g., mire vs. forest) and management practices (e.g., clear‐cutting) determined their spatial variability. Our study advocates the need for integrating terrestrial and aquatic fluxes at the landscape scale based on tall‐tower eddy covariance measurements combined with biomass stock and stream monitoring to develop a holistic understanding of the NLCB of managed boreal forest landscapes and to better evaluate their potential for mitigating climate change.     

Multiple successional pathways in human‐modified tropical landscapes: new insights from forest succession,forest fragmentation and landscape ecology research

Old‐growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human‐modified tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio‐economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land‐use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio‐temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well‐preserved biodiversity‐rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales.     

Aspen lichens in agricultural and forest landscapes: the importance of habitat quality

The amount of aspen Populus tremula , has declined in the boreal forest landscape. This decline is especially marked in young and intermediate stands due to the lack of regeneration. Aspen regeneration is nowadays mainly restricted to abandoned agricultural land. The decrease of aspen is of particular concern as it has more host-specific species than any other boreal tree species. The main question addressed is whether regenerating aspen stands in agricultural habitats can compensate for the deficiency of young stands in the forest. Data on epiphytic macrolichens show that cyanolichens increased, in number and frequency, with stand age in the forest landscape, and that there was a striking difference in species composition between stands in the two landscapes. Lichens with cyanobacterial and green-algal photobionts dominated in the forest and agricultural stands, respectively. Notably, cyanolichens were not found in stands younger than 50 yr in the forest, and stands younger than 100 yr in the agricultural landscape. This difference between the landscapes cannot be explained by stand age, stand size or isolation. Instead, differences in habitat quality, due to differences in the physical environment associated with the presence of conifers in the older forest stands, appear to be involved. We suggest that in order to conserve cyanolichens that are confined to aspen, active management practices have to be adopted that promote the regeneration of aspen in the forest landscape, and the establishment of conifers in areas where aspen regeneration is confined to the agricultural landscape. In addition, until new aspen stands with appropriate physical environments have been established, these measures must be combined with the preservation of existing old-growth stands, which can provide appropriate source populations.