The importance of seed trees in the dioecious conifer Pilgerodendron uviferum for passive restoration of fire disturbed southern bog forests

Biological legacies are important for ecosystem recovery following disturbance as demonstrated by studies in the northern hemisphere. Southern bog forests dominated by the conifer Pilgerodendron uviferum in Northern Patagonia are a typical case of an ecosystem with low resilience to disturbance by fire, which kills most trees and seeds, on which the species depends for regeneration. In this study, we hypothesize that the natural recovery of P. uviferum populations in burned areas is limited by seed availability and this limitation may be exacerbated by the dioecy of the species. Using a multi‐scaled approach, we quantified the seed dissemination potential from P. uviferum seed trees, assessed the suitability of substrates for the germination of seedlings, and finally analysed the spatial distribution of seed trees of the species at the landscape level. Our results indicate that 70 years after a fire on Chiloé Island (43°S), natural regeneration from seed trees can assist the recovery of P. uviferum populations following large‐scale fire disturbance, but their effect is limited at a landscape level owing to a low number of reproductive female trees (0.3 trees ha^?1) and limited seed dispersal (<20 m). In this context, a mixed passive‐active restoration approach that takes into account the spatial pattern and sex of seed trees could be the most effective and efficient option to restore not only P. uviferum forests in North Patagonia, but also other heavily disturbed forests with few remnant seed trees, in particular of dioecious species.     

Shrub influences on seedling performance when restoring the slow‐growing conifer Pilgerodendron uviferum in southern bog forests

Forest restoration is most efficient if it can take advantage of facilitative interactions between established vegetation and planted trees. However, positive and negative interactions have been identified in a number of plant communities. After centuries of anthropogenic fires, forest recovery has been extremely slow in southern bog forests previously dominated by the slow‐growing and vulnerable conifer Pilgerodendron uviferum on Chiloé Island, Chile. Today, the landscape is dominated by secondary shrublands with scattered patches of Sphagnum moss and limited natural tree regeneration. We hypothesized that the retention of secondary shrubs facilitates the early performance of P. uviferum restoration plantings by providing better microsite conditions. To test this hypothesis, we compared the response of seedlings planted on sites prepared at two levels of intervention: after shrubs had been removed or where shrubs were retained. Shrub retention showed a nurse‐plant effect on P. uviferum seedlings 4 years after planting, which resulted in reduced physiological stress (measured as Fv/Fm) for seedlings, as well as reduced browsing. Consequently, the seedlings growing in areas with shrub retention had larger height increment and higher vitality than those in areas where shrubs had been removed. Thus, the more open micro‐site conditions created by shrub removal resulted in generally poorer seedling performance, although seedling mortality—which was low overall (approximately 2–4%)—showed no significant difference between the two levels of intervention. These findings have direct implications for the restoration of slow‐growing conifers that can tolerate extreme wet conditions in highly degraded forests.     

Stand dynamics during a 12-year period in a second-growth stand in a cool temperature forest in northern Japan

In 1981 and 1993, trees over 2 m high were measured and mapped to clarify stand dynamics in two permanent plots of 0.1 ha in a secondary, cool temperate, mixed broadleaf/conifer forest after logging in the Tomakomai Experiment Forest, central Hokkaido, northern Japan. The species could be placed in two groups according to the change in basal area and density, and annual height growth:Quercus mongolica var.grosseserrata, Phellodendron amurense, Prunus sargentii andTilia japonica (Group I: GI) showed episodic regeneration and/or fast height growth;Acer mono, Acer palmatum var.matsumuae andSorbus alnifolia (Group II: GII) showed continuous regeneration and/or slow height growth. The age distribution of stems over 10 cm in d.b.h. suggests synchronous regeneration of GI and GII species. Vertical stratification was promoted during the 12-year period by the difference in annual height growth between the two species groups. Additional tree censuses in both remnant old-growth stands and second-growth stands after large scale blowdowns demonstrated thatQ. mongolica var.grosseserrata is most dominant in stands varying in the stages of development. Disturbance history and successional trends in dominant species implied that stand-devastating disturbances were responsible for the regeneration ofQ. mongolica var.grosseserrata in the study forest.     

Age structure and dynamics of Patagonian beech forests in Torres del Paine National Park,Chile

This study documents the stem size and age-structure in forests dominated by different species of Nothofagus in Torres del Paine National Park (51° S), in the Chilean Patagonian region. We also explored the relationship between the various types of Nothofagus forest and postglacial succession. Pioneer stands on moraine fields 1–2 km of the glacier front are dominated by Nothofagus betuloides and Nothofagus antarctica. Moraines appear to be first colonized by the evergreen N. betuloides, followed within 5–7 years by deciduous N. antarctica. Nothofagus antarctica may replace the former species and develop monospecific stands on glacial valleys. Most trees in the N. antarctica stand studied were older than 40 years and floods may cause a significant mortality of young trees. Recruitment from seed seems to be infrequent. Old-growth stands dominated by deciduous Nothofagus pumilio occupy more stable substrates, and probably represent the last stage of postglacial succession. This long-lived tree species had recorded ages over 200 years. The canopy of N. pumilio forests appears to be a mosaic of even-aged, old-growth patches. We propose that regeneration episodes follow the blowdown of a large portion of the canopy, with long intervals with little or no regeneration. Windstorms may be an important force influencing the regeneration of N. pumilio. Exogenous disturbances, such as floods and windstorms, are an integral part of the forest cycle in the Patagonian region.     

Landscape- vs Gap-level Controls on the Abundance of a Fire-sensitive,Late-successional Tree Species

Tsuga heterophylla (western hemlock), a fire-sensitive, late-successional tree species, is an important component of old-growth forests in the Pacific Northwest, USA. In the Oregon Coast Range, however, T. heterophylla occurs at low densities in or is completely absent from many conifer stands. We used a cellular automata-based simulation model to explore the influences of the fire regime and gap disturbances on T. heterophylla dynamics at a landscape scale. The abundance of T. heterophylla in the simulated landscape was particularly sensitive to variation in fire return interval and fire severity. T. heterophylla was less sensitive to canopy gap return interval, probability of recruitment in canopy gaps, and the probability of rare long-distance dispersal events. Relatively short periods of high fire frequency caused a rapid and persistent decline in the amount of T. heterophylla in the landscape. Based on the simulation results, we propose two new hypotheses. The first is that landscape-level constraints imposed by the frequency, severity, and pattern of wildfires will limit the potential for gap-level processes to affect T. heterophylla regeneration patterns at the landscape scale. The second, related hypothesis is that legacies of cumulative fire effects will have a strong influence on the subsequent development of late-successional forest structure. Received 29 August 2000; accepted 18 October 2001.     

Long‐term dynamics of the long‐lived conifer Libocedrus bidwillii after a volcanic eruption 2000 years ago

Abstract. Question: Following a volcanic eruption of ca. 232 AD, known as the Taupo eruption, the emergent conifer Libocedrus bidwillii expanded on Mt. Hauhungatahi, upwards above the current tree‐line, and downwards into the mixed montane forest. We ask: (1) if current age‐structures at different altitudes support the patterns predicted by the temporal stand replacement model, with cohort senescence and progressively depleting recruitment at ca. 600 year intervals (average cohort age) since the eruption: and (2) if the case history of the population sheds light on the persistence of mixed conifer‐hardwood forests in general. Location: Mt. Hauhungatahi, Tongariro National Park, New Zealand. Methods: The species composition and structure of seven stands covering the altitudinal range of Libocedrus bidwillii, were quantified. Libocedrus trees were cored, and regression equations used to predict ages. Cohorts were identified. Results: Libocedrus densities and basal areas, and the abundance of seedlings and saplings, peaked at different altitudes. At the species’lower limits there has been no recruitment for ca. 550 years, and the angiosperm Weinmannia racemosa has gained dominance. In the tree line and sub‐alpine forest stands, a low level of continuous regeneration has been boosted by periodic cohort recruitment following exogenous disturbances. Conclusions: In the montane zone, the Libocedrus age structure, and its replacement by Weinmannia, are consistent with a model of depleting cohorts separated by ca. 600 years since the Taupo eruption. At higher altitudes more frequent disturbances and reduced competition have allowed Libocedrus persistence. Comparison with other studies suggests long‐term relationships between gymnosperms and angiosperms are mediated by the scale and frequency of disturbance.     

Long-Term Persistence and Fire Resilience of Oak Shrubfields in Dry Conifer Forests of Northern New Mexico

Extensive high-severity fires are creating large shrubfields in many dry conifer forests of the interior western USA, raising concerns about forest-to-shrub conversion. This study evaluates the role of disturbance in shrubfield formation, maintenance and succession in the Jemez Mountains, New Mexico. We compared the environmental conditions of extant Gambel oak (Quercus gambelii) shrubfields with adjoining dry conifer forests and used dendroecological methods to determine the multi-century fire history and successional dynamics of five of the largest shrubfields (76–340 ha). Across the study area, 349 shrubfields (5–368 ha) occur in similar topographic and climate settings as dry conifer forests. This suggests disturbance, rather than other biophysical factors, may explain their origins and persistence. Gambel oak ages and tree-ring fire scars in our sampled shrubfields indicate they historically (1664–1899) burned concurrently with adjoining conifer forests and have persisted for over 115 years in the absence of fire. Aerial imagery from 1935 confirmed almost no change in sampled shrubfield patch sizes or boundaries over the twentieth century. The largest shrubfield we identified is less than 4% the size of the largest conifer-depleted and substantially shrub-dominated area recently formed in the Jemez following extensive high-severity wildfires, indicating considerable departure from historical patterns and processes. Projected hotter droughts and increasingly large high-severity fires could trigger more forest-to-shrub transitions and maintain existing shrubfields, inhibiting conifer forest recovery. Restoration of surface fire regimes and associated historical forest structures likely could reduce the rate and patch size of dry conifer forests being converted to shrubfields.     

The significance of life history strategies in the developmental history of mixed beech (Nothofagus) forests,New Zealand

Size and age structure analysis, dated past disturbances, treefall replacement patterns, and spatial pattern analysis were used to reconstruct the developmental history of two old-growth Nothofagus fusca/N. menziesii stands, South Island, New Zealand. Diameter and height class distributions suggested that N. menziesii was replacing N. fusca, however, stand history reconstruction analysis showed that both species had regenerated intermittently after small-scale disturbances. Although large-scale disturbances such as blowdowns may occasionally generate even-aged stands, gap-phase regeneration maintains the forests in compositional equilibrium. In the absence of other competing tree species and understorey plants the two species appear to coexist by way of different life history strategies, where one species (N. menziesii) has low juvenile mortality and the other (N. fusca) has faster height growth rates and greater longevity and adult survivorship.     

Dynamics of forests with Araucariaceae in the western Pacific

Abstract. Several species of Araucaria and Agathis (Araucariaceae) occur as canopy emergents in rain forests of the western pacific region, often representing major components of total stand biomass. New data from permanent forest plots (and other published work) for three species (Araucaria hunsteinii from New Guinea, A. laubenfelsii from New Caledonia, and Agathis australis from New Zealand) are used to test the validity of the temporal stand replacement model proposed by Ogden (1985) and Ogden & Stewart (1995) to explain the structural and compositional properties of New Zealand rain forests containing the conifer Agathis australis. Here we propose the model as a general one which explains the stand dynamics of rain forests with Araucariaceae across a range of sites and species in the western Pacific. Forest stands representing putative stages in the model were examined for changes through time in species recruitment, growth and survivorship, and stand richness, density and basal area. Support for the model was found on the basis of: 1. Evidence for a phase of massive conifer recruitment following landscape-scale disturbances (e.g. by fire at the Huapai site, New Zealand for Agathis australis); 2. Increasing species richness of angiosperm trees in the pole stage of forest stand development (i.e. as the initial cohort of conifers reach tree size; >10 cm DBH); 3. A high turnover rate for angiosperms (<100 yr), and low turnover for conifers (≥ 100 yr) in the pole stage, but similar turnover rates for both components (50–100 yr) as forests enter the mature to senescent phase for the initial conifer cohort; 4. Very low rates of recruitment for conifers within mature stands, and projected forest compositions which show increasing dominance by angiosperm tree species; 5. A low probability of conifer recruitment in large canopy gaps created by conifer tree falls during the initial cohort senescent phase, which could produce a second generation low density stand in the absence of landscape scale disturbance; 6. Evidence that each of the three species examined required open canopy conditions (canopy openness > 10 %) for successful recruitment. The evidence presented here supports the temporal stand replacement model, but more long-term supporting data are needed, especially for the phase immediately following landscape level disturbance.     

Legacy of Pre-Disturbance Spatial Pattern Determines Early Structural Diversity following Severe Disturbance in Montane Spruce Forests

Background

Severe canopy-removing disturbances are native to many temperate forests and radically alter stand structure, but biotic legacies (surviving elements or patterns) can lend continuity to ecosystem function after such events. Poorly understood is the degree to which the structural complexity of an old-growth forest carries over to the next stand. We asked how pre-disturbance spatial pattern acts as a legacy to influence post-disturbance stand structure, and how this legacy influences the structural diversity within the early-seral stand.

Methods

Two stem-mapped one-hectare forest plots in the Czech Republic experienced a severe bark beetle outbreak, thus providing before-and-after data on spatial patterns in live and dead trees, crown projections, down logs, and herb cover.

Results

Post-disturbance stands were dominated by an advanced regeneration layer present before the disturbance. Both major species, Norway spruce (Picea abies) and rowan (Sorbus aucuparia), were strongly self-aggregated and also clustered to former canopy trees, pre-disturbance snags, stumps and logs, suggesting positive overstory to understory neighbourhood effects. Thus, although the disturbance dramatically reduced the stand’s height profile with ~100% mortality of the canopy layer, the spatial structure of post-disturbance stands still closely reflected the pre-disturbance structure. The former upper tree layer influenced advanced regeneration through microsite and light limitation. Under formerly dense canopies, regeneration density was high but relatively homogeneous in height; while in former small gaps with greater herb cover, regeneration density was lower but with greater heterogeneity in heights.

Conclusion

These findings suggest that pre-disturbance spatial patterns of forests can persist through severe canopy-removing disturbance, and determine the spatial structure of the succeeding stand. Such patterns constitute a subtle but key legacy effect, promoting structural complexity in early-seral forests as well as variable successional pathways and rates. This influence suggests a continuity in spatial ecosystem structure that may well persist through multiple forest generations.     

Forest recovery after swidden cultivation across a 40-year chronosequence in the Atlantic forest of southern Bahia, Brazil

Tree species composition and structure of a 40-year chronosequence of secondary forests was compared with old-growth forests in southern Bahia, Brazil. Twelve stands were randomly selected that represented three age classes: 10, 25, and 40 year old with four replications in each class. All stands selected had been established after abandonment from swidden cultivation and were surrounded by old-growth forests. In every stand, ten 0.01-ha transects were established and all stems (≥5 cm diameter at breast height) were measured and identified. Results were compared with the dataset of two neighboring old-growth sites. Mean diameter, total height, and stand basal area increased with age. Number of trees/ha peaked in 40 year old stands. The results showed that secondary forests in this region take much more than 40 years to recover the structure of old-growth forests. In contrast, species richness recovery was rapid with a continuous accumulation of species with age in secondary forests. Species richness and diversity increased with age as did similarity between secondary stands and old-growth stands. More than half of the species found in the 40 year old stands were shared with the neighboring old-growth forests. However, species richness and diversity were higher in old growth sites.     

Structure and dynamics of subalpine forests in the Wang Lang Natural Reserve, Sichuan, China

Population structure (size, age, spatial patterns) and radial growth patterns are used to analyze regeneration patterns of Abies faxoniana, Betula albosinensis, Betula utilis, Larix potaninii, Picea purpurea, and Sabina saltuaria and reconstruct disturbance history in 8 subalpine forest stands in Wang Lang Natural Reserve, Sichuan, China. In old-growth stands tree regeneration occurs in tree-fall gaps whereby A. faxoniana, Betula sp., P. purpurea, and S. saltuaria persist at stand scales by gap-phase regeneration. Clump sizes of young populations are similar to canopy gap sizes but clumps sizes vary among species. Young Betula patches are larger than those of A. faxoniana suggesting that gap-partitioning by size contributes to species coexistence in mixed stands. Picea purpurea and S. saltuaria are longer lived than A. faxoniana which may compensate for lower recruitment and prevent their replacement by A. faxoniana. Tree regeneration and community structure are also influenced by the understory bamboo Fargesia denudata. Seedlings, saplings, and shrub density all decline with an increase in bamboo cover. Species that regenerate in old-growth forest also regenerate after flooding as do species that establish only on bare substrates (i.e. Larix potaninii, Prunus sp.). Structural and compositional patterns in Wang Lang forests are a reflection of disturbance history, canopy species life history attributes such as dispersal ability, shade tolerance, growth rates, and longevity, and competition of trees and shrubs with understory bamboos.     

The Effects of Land-use History on Soil Properties and Nutrient Dynamics in Northern Hardwood Forests of the Adirondack Mountains

Soil nutrient pools and nitrogen dynamics in old-growth forests were compared with selectively logged stands and stands that were selectively logged and then burned approximately 100 years ago to test the hypothesis that land-use history exerts persistent controls on nutrient capital and nitrogen (N) transformation rates. We provide estimates of net N mineralization and nitrification rates for old-growth forests from the northeastern United States, a region in which few old-growth forests remain and for which few published accounts of mineralization rates exist. At the plot level, no effects of the dominant tree species were observed on any measured soil properties or N-cycling rates. Effects of alternate disturbance histories were detected in soil carbon (C) and N pools. Old-growth forest soils had higher total C (67 Mg·ha^–1) and N capital (3.3 Mg·ha^–1) than that of historically logged then burned soils (C = 50 Mg·ha^–1 and N = Mg·ha^–1), with intermediate values (C = 54 Mg·ha^–1 and N = 2.7 Mg·ha^–1) in the stands that were historically logged. Despite these differences in C and N content, corresponding differences in C–N ratio, net N mineralization rates, and net nitrification rates were not observed. The N concentration in the green foliage of American beech trees (Fagus grandifolia) was also highest from canopy trees growing in old-growth stands (3.0%), followed by logged stands (2.6%), and lowest in the logged/burned stands (2.2%). These data suggest that some legacies of light harvesting on ecosystem processes may be detected nearly 100 years following the disturbance event. These results are discussed in the context of how multiple forest disturbances act in concert to affect forest dynamics.     

Will forest conservation areas protect functionally important diversity of fungi and lichens over time?

Incorporating functional values in biodiversity monitoring systems could add novel perspectives of the status of biodiversity in conservation areas. Stable frequencies of large foliose nitrogen-fixing cyanolichens likely have positive effects on the nitrogen budget of forests and provide food, material and shelter for invertebrates, gastropods and birds. Stable volumes of deadwood and frequencies of associated fungi provide an important supporting function for ecosystem services such as nutrient cycling, carbon storage and soil formation. Based on regional monitoring data from boreal old-growth forest nature reserves and key habitats, we tested for changes in the frequency of various functionally important substrates and species over time. We detected significant reductions in the frequency of indicator cyanolichens occurring on deciduous trees already after 10 years in key habitats, despite non-significant changes in their host substrates. Frequencies of indicator pendulous lichens Alectoria sarmentosa and Bryoria nadvornikiana had also decreased in key habitats, despite overall stable volumes of large conifer host trees. Lichen reductions were more pronounced in the smaller key habitats compared to the larger formally protected nature reserves, likely due to degrading fragmentation and isolation effects. In contrast to these lichens, the average frequencies of old-growth forest indicator fungi decaying coniferous deadwood and common fungi on deciduous trees (Fomes fomentarius) and coniferous trees (Fomitopsis pinicola) remained unchanged. The studied cyanolichens and fruiting fungi generally had similar extinction rates over 10 years, whilst only cyanolichens had substantially lower colonization rates. Amid a severely fragmented landscape, conservation areas seem to struggle in preserving some of the basic old-growth forest values.     

The direct regeneration hypothesis in northern forests

Question: Can the direct regeneration hypothesis (DRH) be used to predict post‐disturbance regeneration after fire, wind disturbance, and clearcutting in northern forests? Do life‐history traits such as regeneration strategy and shade tolerance influence post‐disturbance regeneration success of tree species? Location: Northern forests in North America. Methods: A meta‐analysis was conducted by collecting published data on pre‐ and post‐disturbance stand compositional characteristics in the northern forests. For each tree species, compositional difference (CD) was calculated as the difference between basal area proportions of the post‐ and pre‐disturbance stands, but for post‐disturbance stands <25 years of age, post‐disturbance proportions were calculated based on relative stem density. Results: Species response to disturbances was best explained by regeneration strategy, while disturbance type had no effect on CD. The proportion of broadleaf trees with either strong or weak vegetative reproduction ability increased after all disturbances. Serotinous species had CD values not significantly different from zero after fire, while CD for semi‐serotinous species was negative. The post‐disturbance proportions of non‐serotinous conifers decreased after all forms of disturbance. Conclusions: All disturbances promote broadleaf trees, regardless of regeneration strategy (suckering, sprouting, or seeding). The DRH is supported for conifers with serotinous cones after fire. Fire causes local extinction of non‐serotinous conifers, while wind and clearcutting only decrease the proportion of non‐serotinous conifers because of partial survival of seed sources and advanced regeneration. This study suggests that increasing stand‐replacing disturbances associated with global climate change will promote broadleaf trees in northern forests.     

Species diversity of longicorn beetles in humid warm-temperate forests: the impact of forest management practices on old-growth forest species in southwestern Japan

In the humid warm-temperate zone of southwestern Japan, old-growthforests have been seriously fragmented to small remnants due to traditionalagriculture and coppicing as well as recent rapid plantation with conifers.Assemblages of longicorn beetles (Coleoptera: Disteniidae and Cerambycidae) werecompared among old-growth forests, second-growth forests and conifer plantationsusing collision traps baited with chemical attractants. Species richness oflongicorn beetles was poorer in second-growth forests and conifer plantationsthan in old-growth forests. It was proved by multidimensional scaling(MDS) that the beetle assemblages of old-growth forests were distinct from thoseof conifer plantations, while those of second-growth forests were intermediatebetween them. Further analysis showed that a number of species, including manyPidonia spp., were specific to or closely associated withold-growth forests, and the results were largely supported by the indicatorvalue (IndVal) approach. It is likely that many of such old-growth forestspecies in the larval and pupal stages require large broad-leaved trees standingor fallen with thick bark. At the same time, the flower-visiting adults wouldplay an important role in pollinating various herbaceous and woody plants.Regional forest management for the conservation of insect biodiversity is alsodiscussed.     

Old trees as a key source of epiphytic lichen persistence and spatial distribution in mountain Norway spruce forests

Habitat loss and fragmentation can negatively impact the persistence of dispersal-limited lichen species with narrow niches. Rapid change in microclimate due to canopy dieback exposes species to additional stressors that may limit their capacity to survive and colonize. We studied the importance of old trees as micro-refuges and microclimate stability in maintaining lichen survival and diversity. The study was situated in mountain Norway spruce (Picea abies) forests of the Gorgany Mountains of the Ukrainian Carpathian mountain belt. Lichens were collected on 13 circular study plots (1000 m²). Dendrochronological methods were used to reconstruct age structure and maximum disturbance event history. A linear mixed effects model and general additive models were used to explain patterns and variability of lichens based on stand age and disturbance history for each plot. Tree age was the strongest variable influencing lichen diversity and composition. Recent (<80 years ago) severely disturbed plots were colonized only by the most common species, however, old trees (>200 years old) that survived the disturbances served as microrefuges for the habitat-specialized and/or dispersal limited species, thus epiphytic lichen biodiversity was markedly higher on those plots in comparison to plots without any old trees. Most species were able to survive microclimatic change after disturbances, or recolonize disturbed patches from surrounding old-growth forests. We concluded that the survival of old trees after disturbances could maintain and/or recover large portions of epiphytic lichen biodiversity even in altered microclimates.     

Long-term persistence of aspen – a key host for many threatened species – is endangered in old-growth conservation areas in Finland

Large, dead and dying European aspens (Populus tremula L.) host many threatened species in Fennoscandian boreal forests. Large aspen trees have mostly disappeared and are being harvested from the managed forests that cover 95% of the forest area in Finland. Due to the small area protected (4.1%), the aspen-associated species may encounter major difficulties in the protected areas if aspen trees disappear due to natural forest succession. The availability of aspens was assessed in the old-growth conservation area network in eastern Finland. We mapped all the living and dead aspens in 15 protected old-growth forests. The total number of counted trees was 32 903 individuals. Current amounts of living (2.7 m³/ha) and especially dead aspens (2.8 m³/ha) in the protected areas were higher than in the surrounding managed forests (1.1 and 0.1 m³/ha for living and dead trees, respectively). However, while saplings (dbh<5 cm) occur in most of the areas (12 individuals/ha on average) they survive poorly and young aspen cohorts (5 cm <dbh<15 cm) are lacking or are very rare. The most likely reason for the poor sapling survival is high browsing pressure by the mammalian herbivores, especially the moose. The moose population has increased many times in Finland during the past decades. The poor regeneration of aspens implies that the value of the old-growth conservation areas for aspen-associated species will face a serious bottleneck within a few decades when the currently middle-aged tree cohorts disappear. If the current high browsing pressure and lack of natural disturbances continue the obligatory aspen-associated species may disappear both locally and regionally from the network of the protected areas.     

Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan

To evaluate the effects of canopy gaps and forest floor microsites (soil, fallen logs, root-mounds, buttresses and stumps) on regeneration of subalpine forests, the gap regeneration and seedling occurrence of conifers (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis and Tsuga diversifolia) were studied in two stands of a subalpine old-growth forest, central Japan. The percentage of gap area to total surveyed area was 11.2–11.3% in the stands. Gap regeneration was not common for P. jezoensis var. hondoensis and T. diversifolia. In contrast, gap regeneration by advanced regeneration was relatively common for Abies. Seedling occurrence of P. jezoensis var. hondoensis and T. diversifolia was restricted on elevated surfaces such as stumps and root-mounds, while Abies seedlings could occur on soil as well as on elevated surfaces. Rotten stumps were the most favorable microsites for conifer seedling occurrence, which covered small area in the forest floor. Although canopy gaps were not always favorable for seedling occurrence, all conifer seedlings were larger under canopy gaps than under closed canopy. Canopy gaps and forest floor microsites clearly affected seedling occurrence and growth of conifers. This suggests that regeneration of conifers is related to the difference of growth advantage under canopy gaps and favorable microsites for seedling occurrence.     

Old rural parks support higher biodiversity than forest remnants

One of the main challenges in biodiversity conservation is to curb a further degradation and loss of high-quality habitats. In agricultural matrix landscapes, the detection of alternative habitats for habitat specialists may be a solution. Historic old parks or landscape gardens around manor houses and castles are cultural heritage of nobles, but their value in harbouring biodiversity is poorly acknowledged. Therefore we evaluated the potential of old rural parks to serve as a habitat for nemoral forest species. We recorded stand structure and the presence of forest biodiversity indicators in 74 closed-canopy stands of historic parks and compared them with 93 neighbouring mature forest remnants on ancient forest land. We estimated the importance of stand structure in relation to habitat type on biodiversity indicators. Finally we suggest single-value indicator-complexes for the cost-efficient assessment of the conservation value of forests and forest-like habitats. Park stands outclassed reference forests in several individual structural characteristics, and in combined indicators of habitat quality and biodiversity. Forests had higher estimates for the combined indicator of dead wood, but large-diameter dead wood types were more abundant in parks. Woodpeckers, several old-growth indicator epiphytes and forest herbs had successfully become established in planted forest-like park fragments. Old rural parks resemble high-conservation-value forests more than the best preserved contemporary forest remnants. After the century needed to overcome immigration delay, old parks do provide a refugium for temperate deciduous forest species. Consequently, biodiversity-targeted management should retain and enhance old-growth attributes in forests and on the peripheries of parks: e.g. preserving old trees to provide service for epiphytes, hollow trees and an understorey mosaic for birds and bats; dead wood elements for saproxylic insects and fungi; limited mowing frequency and increased cutting height for forest herbs. Forestry should enhance the recovery of mixed deciduous stands and avoid conifer plantations.