Planning timber harvest of residual forest stands without compromising bird and small mammal communities in boreal landscapes

Numerous efforts have been invested in designing and configuring residual forest stands in Canadian boreal forest to preserve their overall biodiversity. Now that several landscapes have been partially logged, the next issue in forest management involves the planning of residual forest stand harvesting without compromising wildlife populations. Residual stands can be cut when adjacent regeneration reaches 3 m in height according to current regulations in several Canadian provinces (e.g., Québec, Ontario, Alberta, and British-Columbia). However, little is known on whether such regenerating habitat (RE-3m) can maintain wildlife communities similar to those found in unharvested mature forest (CO). We estimated the relative abundance of small mammals and forest birds in RE-3m and CO habitats and characterized landscape and stand structures. These variables were then compared between the two contrasting successional stages and were used to build habitat use models (HUMs) for 21 species. CO and RE-3m differed with regard to several landscape characteristics and stand structure variables as a result of logging. Snowshoe Hare, Northern Flicker, Alder Flycatcher, Ruby-crowned Kinglet, White-throated Sparrow and Magnolia Warbler were more abundant in RE-3m than CO, while Red-backed Vole, Brown Creeper and Golden-crowned Kinglet exhibited lower abundances in RE-3m. No significant differences in abundance were observed for the 12 other species. Species HUMs were highly significant and explained between 64.3 and 99.1% of the total variability in abundance. Following variance partitioning, stand structure variables accounted for most of the explained variability (54.2%) while landscape characteristics accounted for only 28.7%. No difference in species richness was observed but community evenness was greater in CO than RE-3m. Our results suggest that current regulations may threaten the maintenance of 3 out of 21 censused species for which abundances were significantly lower in regenerating 3 m tall stands. As stand structure explained a large amount of variability in abundance, it should be considered during timber harvest planning in both mature and regenerating stands. Until we know more on whether the current regulations are suitable for maintaining overall biodiversity, our results suggest that some mature forest stands should be maintained within managed landscapes for a complete logging rotation period.     

Effects of artificial afforestation and successional stage on a lowland forest bird community in southern China

To examine the relationship between forest succession after severe logging forestry practices and the composition of avian communities, we investigated how forest bird composition and guild structure change as a function of structural properties along a successional gradient, including a climax mature forest (>400 years), a rehabilitated mixed forest (50-70 years), and a disturbed Masson pine forest (70 years) of the Dinghushan Nature Reserve, Guangdong Province, China. Of a total of 51 resident species recorded, mixed forests hosted the highest numbers of individuals and species, reflecting the high species richness of both forest and non-forest species. For forest-dependent species, however, mature stands had the highest observed and estimated species richness. Of 6 habitat-use guilds identified, vertical-profile generalists and understory-birds formed the two dominant guilds, accounting for 54.0% and 38.7% of all individuals respectively. The results of canonical correspondence analysis (CCA) clearly showed that most forest-dependent species were associated with high proportions of native canopy cover and the mean density of dead trees and large trees, which are characteristic of old-growth mature forests (horizontal heterogeneity) at stand level. Accordingly, conservation efforts should focus on the specialized requirements of the most habitat-restricted species in the future, especially for understory insectivores (Babblers) and large-tree users in mature subtropical monsoon forests of southern China. Moreover, since regenerating mixed forests are very similar to mature forests in both vegetation structure and bird community composition, we recommend that logging cycles (>50 years) be increased to a minimum of 50 years in southern China, so that a balance between economic and ecological interest can be reestablished.     

Stand and landscape-level factors related to bird assemblages in exotic pine plantations: Implications for forest management

Plantations cover a substantial amount of Earth's terrestrial surface and this area is expected to increase dramatically in the coming decades. Pinus plantations make up approximately 32% of the global plantation estate. They are primarily managed for wood production, but have some capacity to support native fauna. This capacity likely varies with plantation management. We examined changes in the richness and frequency of occurrence of bird species at 32 plots within a Pinus radiata plantation (a management unit comprising multiple Pinus stands) in south-eastern Australia. Plots were stratified by distance to native forest, stand age class and thinning regime. We also assessed the landscape context of each plot to determine relationships between bird assemblages and stand and landscape-level factors. Bird species richness was significantly higher at plots ≥300 m from native forest and in mature (∼20 years since planting) and old (∼27 years since planting) thinned pine stands. We were able to separate the often confounding effects of stand age and thinning regime by including old stands that had never been thinned. These stands had significantly fewer species than thinned stands suggesting thinning regime, not age is a key factor to improving the capacity of pine plantations to support native species (although an age × thinning interaction may influence this result). At the landscape level, species richness increased in pine stands when they were closer to native riparian vegetation. There were no significant differences in species composition across plots. Our study indicates the importance of stand thinning and retention of native riparian vegetation in improving the value of pine plantations for the conservation of native fauna.     

Bird communities in hemiboreal old-growth forests: The roles of food supply, stand structure, and site type

Key factors causing the difference of wildlife populations in natural and managed forests are an important field of ecosystem and biodiversity research. To explore the factors contributing to bird-community features in the poorly studied European natural hemiboreal forests, we carried out a comparative study in old-growth and mature stands of five site types in Estonia. The mature stands were of clear-cut origin and managed for timber production. Old-growth hosted both more diverse and more abundant bird communities than mature stands, which does not support the putative ‘old-growth syndrome’ (high diversity at a low density) described previously in temperate Europe. Site-type specificity of bird communities was also more pronounced in old-growth, indicating a timber-harvesting induced process of biotic homogenization. In particular, natural swamp forests had characteristic bird species and those communities may be additionally sensitive to artificial drainage. In terms of invertebrate food supply, the availability of snails, rather than of insects, was related to bird-community characteristics; however, the influence of snails was due to one snail-poor forest type (Vaccinium type pine stands), not management. The abundance of coarse woody debris was the main structural feature affecting bird communities; tree-size variation was additionally important for species richness. A significant unexplained ‘old-growth’ effect remained even after the variables describing food supply and stand structure were taken into account. Our results imply the distinct importance of old-growth of different site types for hemiboreal bird communities. However, we did not obtain any evidence of different key factors structuring the bird communities in old-growth and mature stands.     

The effects of forest structure on the risk of wind damage at a landscape level in a boreal forest ecosystem

? The aim of this work was to analyze how the forest structure affects the risk of wind damage at the landscape level in a boreal forest.

? This was done by employing: (i) Monte Carlo simulation technique for generating landscapes with different age class distributions, proportions of open areas (gaps), and tree species composition; and (ii) a mechanistic wind damage model, HWIND, for predicting the critical wind speeds at downwind stand edges of open areas (gaps) for risk consideration. The level of risk of wind damage observed at the landscape level was significantly affected by the presence of gaps and old stands. Even a slight increase in the proportion of gap areas or older stands had a significant impact on the total length of edges at risk. As a comparison, variation in species composition (Scots pine and/or Norway spruce) had much smaller impact on the risk of damage.

? In conclusion, the effects of forest structure on the risk of wind damage should especially be considered by forest managers in day-to-day forest planning in order to reduce the risk of wind damage both at the stand and landscape level.

     

Silviculture as a disturbance regime: the effects of clear-cutting,planting and thinning on polypore communities in mixed forests

The structure of modern forest landscapes is profoundly affected by human-caused disturbances, particularly forest management; however, the effects and prospects of individual silvicultural techniques are insufficiently understood. This study distinguishes the effects of clear-cutting, planting and thinning on species richness and community composition of polypore fungi. In 2008–2009, 181 forest compartments (ranging from naturally regenerated deciduous stands to planted Picea abies stands and 0–137 years post clear-cutting) were explored in a hemiboreal landscape subjected to even-aged management. Altogether 104 polypore species were recorded. For species richness, time since clear-cutting was the most influential factor at both stand and landscape scales, followed by thinning. Clear-cuts had distinct polypore communities (including several red-listed species) whose species richness declined in time. Following 20 years post clear-cutting, species richness started to increase along different community–composition pathways determined by regeneration type. The communities developed after planting were moderately species rich at stand scale but homogeneous over larger areas. Thus, at landscape scale, mature unmanaged naturally regenerated stands hosted most species; thinning reduced species richness by approximately 15%, and among thinned stands, planted areas had a further 9–22% fewer species than naturally regenerated areas. In such variably managed landscape, silviculture appeared to create particularly distinct communities in young stands on nutrient-rich soils, which naturally provide polypores with a rich supply of small deciduous snags absent from stands artificially planted with P. abies and intensively thinned.     

Influence of post-harvest silviculture on understory vegetation: Implications for forage in a multi-ungulate system

Natural disturbance emulation has emerged as a key management approach to maintaining biodiversity in logged boreal forests. Forest managers’ success in emulating understory forest ecosystem functions, e.g., for the provision of habitat even for large mammals, has not been tested due, in many cases, to incomplete records of silviculture. We examined regenerating areas of previously conifer-dominated forests in northwestern Ontario, Canada, 10 and 30 years after logging and 10 and 30 years after fire to test if understory development and moose (Alces alces) forage abundance differed between the two disturbance types and artificial or natural regeneration approaches. In addition, we counted moose pellet groups as a measure of moose use of the region. Specific treatments included: (1) naturally regenerating, fire-origin forests, (2) post-harvest, regenerating forests with natural establishment of trees, and (3) post-harvest, regenerating forests with mechanical or chemical site preparation and planting and/or herbicide spraying. We hypothesized that the understory in post-harvest stands would support higher forage availability for moose compared to similarly-aged, fire-origin stands. Abundance of hardwoods, shrubs, and herbaceous plants was greater in naturally-regenerated post-harvest stands than in fire-origin and artificially regenerated post-harvest stands at both 10 and 30 years post-disturbance. However, post-harvest, naturally regenerating stands were not significantly associated with higher moose use, rather evidence of moose use increased as a function of the amount of naturally regenerating logged forest in the surrounding landscape. This study suggests that, relative to fire, the intensity of post-harvest silviculture influences habitat suitability for moose. The effect likely cascades to other ungulates, such as woodland caribou (Rangifer tarandus caribou), and vegetation management needs to be considered at scales greater than the stand level in order to achieve habitat management for large mammals.     

Variation in post-wildfire regeneration of boreal mixedwood forests: underlying factors and implications for natural disturbance-based management

To test the direct regeneration hypothesis and support natural disturbance-based forest management we characterized the structure and composition of boreal mixedwood forests regenerating after large wildfires and examined the influence of pre-fire stand composition and post-fire competing vegetation. In stands which had been deciduous (Populus sp.)-dominated, conifer (white spruce)-dominated, or mixed pre-fire we measured regeneration stocking (presence in 10 m² plots), density and height 10–20 years post-burn in five wildfires in Alberta, Canada. Most plots regenerated to the deciduous or mixed stocking types; plots in the older fire and in stands that were pure conifer pre-fire had higher amounts of conifer regeneration. Surprisingly, regeneration in pre-fire ‘pure’ white spruce stands was most often to pine, although these had not been recorded in the pre-fire inventory. Pre-fire deciduous stands were the most resilient in that poplar species dominated their post-fire regeneration in terms of stocking, density and height. These stands also had the highest diversity of regenerating tree species and the most unstocked plots. High grass cover negatively affected regeneration density of both deciduous and conifer trees. Our results demonstrate the natural occurrence of regeneration gaps, pre- to post-fire changes in forest composition, and high variation in post-fire regeneration composition. These should be taken into consideration when developing goals for post-harvest regeneration mimicking natural disturbance.     

Weak response of bryophyte assemblages to second commercial thinning in boreal spruce forest of south-central Sweden

Forest management regimes dictate the composition and structure of large parts of the Fennoscandian boreal forest region. The understanding of thinning impacts on understorey plant communities is considerably lower compared to the effects of final felling. We studied the response of bryophyte assemblages to second commercial thinning in spruce-dominated stands sampled to be representative of the dominant forest type of south-central Sweden. Eighteen stands were analysed 6–12 years after second commercial thinning, and were compared with six stands commercially thinned only once, at two spatial scales (0.1 and 0.01?ha), with stand ages varying between 52 and 66 years. We found few strong differences in bryophyte species richness and no differences in bryophyte species composition between stands thinned once or twice. More pioneer species were found in stands thinned only once, a result that needs further study. Species richness declined significantly with time since thinning on both investigated spatial scales. Canopy cover significantly affected the composition of forest floor species. Thinning effects over longer time periods need further study, and also effects in different forest types along productivity gradients. For nature conservation, studies targeted towards thinning response of uncommon species connected to specific habitats and substrates will be essential.     

Effect of selective logging on stand structure and tree species diversity in a subtropical evergreen broad-leaved forest

?Context

Selective logging followed by natural regeneration is rarely employed for restocking subtropical evergreen broad-leaved forests in East Asia compared with the use of clear-cutting.

?Aims

To clarify the succession of these forests, the effects of selective logging on stand structure, species diversity, and community similarity were studied in a mature and regenerating forest in Okinawa, Japan.

?Methods

Four study plots were established, and trees ≥1.2 m height were identified by species name, tree height, and diameter at breast height.

?Results

The results showed that the species composition of regenerating forest was similar to mature forest; however, the former had a greater species density and Shannon–Wiener index than the latter. Castanopsis sieboldii and Distylium racemosum, the predominant trees in the mature forest, continued to dominate the regenerating forest, with a broad layer distribution. High Sørensen and Jaccard community similarity indices for mature and regenerating forest indicated that the regeneration occurred in a progressive succession.

?Conclusion

The similar species composition and stand structure for both mature and regenerating forest, and the higher species diversity for the latter, provided no evidence of forest degeneration and suggested that the regenerating forest may develop into a stand similar to preselective logging forest.     

Predicting breeding bird occurrence by stand- and microhabitat-scale features in even-aged stands in the Central Appalachians

Spatial scale is an important consideration when managing forest wildlife habitat, and models can be used to improve our understanding of these habitats at relevant scales. Our objectives were to determine whether stand- or microhabitat-scale variables better predicted bird metrics (diversity, species presence, and abundance) and to examine breeding bird response to clearcut size and age in a highly forested landscape. In 2004-2007, vegetation data were collected from 62 even-aged stands that were 3.6-34.6 ha in size and harvested in 1963-1990 on the Monongahela National Forest, WV, USA. In 2005-2007, we also surveyed birds at vegetation plots. We used classification and regression trees to model breeding bird habitat use with a suite of stand and microhabitat variables. Among stand variables, elevation, stand age, and stand size were most commonly retained as important variables in guild and species models. Among microhabitat variables, medium-sized tree density and tree species diversity most commonly predicted bird presence or abundance. Early successional and generalist bird presence, abundance, and diversity were better predicted by microhabitat variables than stand variables. Thus, more intensive field sampling may be required to predict habitat use for these species, and management may be needed at a finer scale. Conversely, stand-level variables had greater utility in predicting late-successional species occurrence and abundance; thus management decisions and modeling at this scale may be suitable in areas with a uniform landscape, such as our study area. Our study suggests that late-successional breeding bird diversity can be maximized long-term by including harvests >10 ha in size into our study area and by increasing tree diversity. Some harvesting will need to be incorporated regularly, because after 15 years, the study stands did not provide habitat for most early successional breeding specialists.     

Effect of stand age on bird communities in late-successional Macedonian pine forests in Bulgaria

To gain insight into the question of which vegetation characteristics have the most influence on avian assemblages in late-successional forests, the habitat preferences of bird-guilds in old-growth endemic forests of Macedonian pine were studied over 3 years in the Pirin National Park, Bulgaria. Bird–habitat relationships were investigated by comparing vegetation characteristics, and bird species richness, diversity, abundance, and guild structure of birds (determined according to food type, foraging and nesting sites) between mature (60–100 years old) and over-mature (>120 years old) Macedonian pine forest stands. Studied forest age-classes differed mainly by the density, height and diameter of trees, and the amount of dead wood. The first one of these parameters decreased and the latter two parameters increased with the forest succession. The difference in the vegetation structure affected the abundance of bird-guilds and thus, the overall bird abundance and the structure of avian assemblages within Macedonian pine forests. There was no significant difference in bird diversity among studied forest age-classes, but the overall bird abundance increased with forest maturation. Analyzed by study plots, species richness was higher in over-mature forests, but at cluster level, there was no significant difference between mature and over-mature forest age-classes. Half of the studied (insectivorous, hole- and ground-nesters, bark- and canopy-foraging bird species) guilds were more abundant in over-mature forests, while there was no bird-guild exhibiting a preference for mature forest stands. The abundances of bird-guilds were correlated with tree height, diameter at breast height and the amount of dead wood between the studied forest age-classes and this might explain their preferences for over-mature pine forests. Therefore, for future sustainable management of these endemic forests and the conservation of their avifauna, efforts should focus on protecting the remaining native old-growth forest stands and the importance of the structure of Macedonian pine forests on their bird assemblages should be considered in forestry practices.     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.     

Ectomycorrhizal fungal communities of Quercus liaotungensis along different successional stands on the Loess Plateau,China

Although Quercus liaotungensis forest is an important component of Loess Plateau forest ecosystems, little is known about the ectomycorrhizal fungal (EMF) symbiosis of Q. liaotungensis in this ecosystem. Here we investigated EMF communities of Q. liaotungensis along three successional stands on the central Loess Plateau (Yan’an, China): a seedling stand (1 to 3-year-old seedlings under a Populus davidiana forest stand), a young tree stand (20 to 30-year-old Q. liaotungensis forest stand), and a mature tree stand (50 to 70-year-old Q. liaotungensis forest stand). In each stand, the Q. liaotungensis roots were sampled, then EMF communities were examined on the basis of EMF morphotyping, PCR–RFLP, and DNA sequencing. In total, 70 EMF species were observed, and the total richness of estimated species exceeded 100 EMF species. The EMF community was composed of a few common taxa (approx. 7 %) and many rare taxa (approx. 61 %). EMF richness in young and mature trees was higher than in seedlings. EMF communities of young and mature trees were more similar than those of seedlings and trees. Nonmetric multidimensional scaling ordinations of the EMF community revealed more separation among Q. liaotungensis of different age. This study provides preliminary insight into the taxonomic diversity of EMF of Q. liaotungensis along different successional stands on the Loess Plateau.     

Stand specific occurrence of coarse woody debris in a managed boreal forest landscape in central Sweden

Forests long subjected to management for timber production contain only a fraction of the volumes of coarse woody debris (CWD) found in pristine forests. This is a threat to many organisms that depend on CWD. Forest management practices have been altered to achieve an increase in the amounts of CWD. Few studies have attempted to analyze the occurrence of CWD at the landscape level. We studied the occurrence of CWD in stands of different ages and management background in a boreal forest landscape in central Sweden. Volume of CWD in unmanaged stands (nature reserves and set-asides) was twice that in managed stands. The composition of CWD was influenced by stand age and management regime. Standing CWD was more common in unmanaged stands than in managed stands. Pine CWD was particularly prevalent in young forest stands (8–59 years of age). Bark-covered CWD was most common on deciduous and spruce wood and uncommon on pine. Bark area in young forest stands was almost 10 times lower than that in other managed stands. Using the age distribution of stands, we estimated the volume and bark area of CWD in the landscape. Recent clear-cuts harvested in accordance with new management guidelines contained more early decay CWD per ha than old managed stands. Young forests covered over half the landscape and had significantly lower volumes of spruce and deciduous CWD compared with other stands. The consequences of these results for biodiversity-oriented forest management are discussed.     

Simulated cavity tree dynamics under alternative timber harvest regimes

We modeled cavity tree abundance on a landscape as a function of forest stand age classes and as a function of aggregate stand size classes. We explored the impact of five timber harvest regimes on cavity tree abundance on a 3261 ha landscape in southeast Missouri, USA, by linking the stand level cavity tree distribution model to the landscape age structure simulated by the LANDIS model. Over 100 years, mean cavity tree density increased constantly under all timber harvest regimes except for even-aged intensive management. This was due in large part to the continued maturation of the numerous stands that were >70 years old at the start of the simulations. However, compared to the no harvest (control) regime, the uneven-aged, the mixed, the even-aged long rotation, and the even-aged intensive harvest regimes reduced the cavity tree density by 9–11, 11–13, 15–18, and 28–34%, respectively, as more old stands were cut. Forest managers and planners can use this information to evaluate the practical consequences of alternative timber harvest regimes and consider the need for activities such as cavity tree retention.     

The relationship between shelterwood cuts and crown thinnings and the abundance and distribution of birds in a southern New England forest

Forest regeneration methods such as shelterwood treatments have been shown to substantially increase the diversity of bird species, specifically of species that prefer early seral forests, now rare in the eastern United States. Stand improvement techniques such as thinnings have also been found to increase avian diversity under some conditions. A sustainably managed forest, however, must simultaneously apply regenerative harvesting with stand improvements, and the effect of such treatment combinations on bird community composition is not clear. We compared bird distribution and abundance on shelterwood cuts, crown thinnings, and unmanaged stands at the Yale Myers Forest, a large privately owned and actively managed forest in southern New England. Bird abundance and species diversity was highest in shelterwood cuts and lowest in unmanaged forest, with thinnings being intermediary. Different suites of species inhabited the three treatments, with 18 of 49 common species differing significantly in their abundances between treatments. Characteristics of the vegetation that were directly influenced by silvicultural intervention, including canopy openness, seedling regeneration and vertical structural diversity, appeared to be the dominant drivers of bird abundance. The abundances of some species or groups of species were correlated with the number of trees retained in the implementation of the forestry practices. In conjunction with the conservation of a variety of mature forest habitats, regenerative cuts and stand improvement techniques can be used together to sustain a diverse assemblage of bird species.     

Effects of stand density on ecosystem properties of subalpine forests in the southern Rocky Mountains, USA

? Mixed coniferous, subalpine forest communities in the Rocky Mountains are historically dense and have experienced infrequent, high-severity fire. However, many of these high-elevation stands are thinned for a number of perceived benefits.

? We explored the effects of forest stand density on ecosystem properties in subalpine forests in Colorado, USA, 17–18 y after forests were managed for timber.

? Forest structure significantly altered the composition and chemical signature of plant communities. Previously managed stands contained lower density of overstory trees and higher ground cover compared to paired reference stands. Foliar phenolic concentration of several species was negatively related to basal area of overstory trees. Furthermore, reductions in stand density increased total foliar phenolic:nitrogen ratios in some species, suggesting that gap formation may drive long-term changes in litter quality. Despite significant changes in forest structure, reductions in stand density did not leave a strong legacy in surface soil properties, likely due to the integrity of soil organic matter reserves.

? Changes in forest structure associated with past management has left a long-term impact on plant communities but has only subtly altered soil nutrient cycling, possibly due to trade offs between litter decomposability and microclimate associated with reductions in canopy cover.

     

Are gullies best for biodiversity? An empirical examination of Australian wet forest types

A relatively common observation in forest environments has been that gullies support higher species richness and individual abundance than elsewhere in the landscape. We completed a detailed case study of birds to contrast species richness and assemblage composition between gullies and other parts of the topography of landscapes in three closely related and spatially adjacent wet ash forest types – those dominated by Mountain Ash (Eucalyptus regnans), Alpine Ash (E. delegatensis) or Shining Gum (E. nitens) – in the Central Highlands of Victoria, south-eastern Australia. We also quantified the influence of a wide range of other measures of stand structure and plant species composition on the bird assemblage and on individual bird species.