Linking substrate and habitat requirements of wood‐inhabiting fungi to their regional extinction vulnerability

Loss of old‐growth forests and greatly reduced volumes of coarse dead wood in managed forests are the main reasons for the decline of many wood‐inhabiting species in Europe and elsewhere. To assess the habitat requirements and extinction vulnerability of 13 polypore species associated mainly with spruce, their occurrences were recorded on 96 521 dead‐wood objects in 331 stands along a regional gradient of forest utilization history across southern‐middle boreal Finland. The substrates studied included a variety of tree species and dead‐wood qualities investigated in both unmanaged and managed stands at different successional stages. Hierarchical logistic regression models were constructed to analyze the relationships between the occurrence probability of individual species and variables at the substrate, stand and regional scales. The substrate preferences of the polypore species studied overlapped, since most of them favored large‐diameter spruce logs in mid‐decay stages. However, only a few species were restricted to this substrate. Other species were able to use a wider range of host tree species and qualities of dead wood, including man‐made substrates that are abundant in managed forests (logging residues and stumps). Species confined to logs had a significantly lower occurrence probability in regions with the longest and most intensive forest use history. Species less specialized in their resource use showed no decline or the opposite trend. Loss of threatened species is likely if the preservation of old‐growth forests is not combined with conservation measures in managed forests. Increasing extraction of logging residues and stumps for biofuel may cause non‐threatened species to decline by reducing substrate qualities utilized by them. The hierarchical models predicted a considerable part of the variation in Species' occurrence probabilities, and therefore provide powerful tools for setting quantitative targets for management.     

Structure,dynamics and dendroecology of an old‐growth Fagus forest in the Apennines

Abstract. Question: Which are the structural attributes and the history of old‐growth Fagus forest in Mediterranean montane environments? What are the processes underlying their structural organization? Are these forests stable in time and how does spatial scale affect our assessment of stability? How do these forests compare to other temperate deciduous old‐growth forests? Location: 1600–1850 m a.s.l., Fagus forest near the tree line, central Apennines, Italy. Methods: An old‐growth Fagus forest was studied following historical, structural and dendroecological approaches. History of forest cover changes was analysed using aerial photographs taken in 1945, 1954, 1985 and 1994. The structural analysis was carried out in the primary old‐growth portion of the forest using 18 circular and two rectangular plots. Dendroecological analyses were conducted on 32 dominant or co‐dominant trees. Results: These primary old‐growth Fagus remnants consist of four patches that escaped logging after World War II. Both living and dead tree components are within the range of structural attributes recognized for old‐growth in temperate biomes. Dendroecological analyses revealed the roles of disturbance, competition and climate in structuring the forest. We also identified a persistent Fagus community in which gap‐phase regeneration has led to a mono‐specific multi‐aged stand at spatial scales of a few hectares, characterized by a rotated‐sigmoid diameter distribution. Conclusion: Even at the relatively small spatial scale of this study, high‐elevation Apennine Fagus forests can maintain structural characteristics consistent with those of old‐growth temperate forests. These results are important for managing old‐growth forests in the Mediterranean montane biome.     

Biodiversity conservation in old-growth boreal forest: black spruce and balsam fir snags harbour distinct assemblages of saproxylic beetles

Biodiversity conservation of forest ecosystems strongly relies on effective dead wood management. However, the responses of saproxylic communities to variations in dead wood characteristics remains poorly documented, a lack of knowledge that may impede the development of efficient management strategies. We established the relationship between saproxylic beetles—at the species and community levels—and attributes of black spruce and balsam fir in old-growth boreal forests. The relationship was first evaluated for individual snag bole segments, and then for forest stands. A total of 168 bole sections were collected in summer 2006 along a compositional gradient ranging from black spruce-dominated stands to balsam fir-dominated ones, in a boreal forest dominated by >90-year-old stands. A total of 16,804 beetles belonging to 47 species emerged from bole segments, with 21% of the species being found exclusively in black spruce snags and 36% exclusively in balsam fir snags. Black spruce and balsam fir snags thus contributed differently to forest biodiversity by being inhabited by different saproxylic communities. Wood density was an important attribute in the host-use patterns for several species of saproxylic beetles, but no relationship was found between snag availability within stands and abundance of beetles strongly linked to either black spruce or balsam fir. Our study outlines the relative contribution of tree compositional diversity to saproxylic species, while highlighting the contribution of black spruce and balsam fir to animal diversity in old-growth boreal forests.     

Present and past old‐growth forests of the Lake Tahoe Basin,Sierra Nevada,US

Abstract. We described 38 relictual old‐growth stands – with data on the mortality, regeneration, floristic richness, fuel load and disease incidence in our study area in the Tahoe Basin of California and Nevada. The stands are within the lower and upper montane zones (1900–2400 m a.s.l.) and they are rare, occupying < 2% of the land in the Basin's watershed. Correlation matrices and ANOVAs of forest types and conifer species with environmental gradients revealed significant relationships with elevation, distance east of the Sierran crest, slope aspect, annual precipitation, date of complete snow melt, litter depth and degree of soil profile development. Pathogens, parasites and wood‐boring insects were present on 23% of living trees; 16% of all trees were dead. We compared these stands to a reconstruction of pre‐contact Basin forests and to ecologically analogous old‐growth forests of Baja California that have never experienced fire suppression management. Currently, overstorey trees (> 180 yr old) in the Basin stands have ca. 33% cover, 54 m².ha^‐1 basal area and 107 individuals.ha^‐1, values very similar to reconstructions of pre‐contact Basin forests and to modern Baja California forests. Understorey trees (60–180 yr old), however, are several times more dense than historic levels and species composition is strongly dominated by A. concolor, regardless of the overstorey composition. The ratio of Pinus: Abies has increased – and the age structure of extant stands predicts that it will continue to increase – from approximately 1:1 in pre‐contact time to 1:7 within the next century. Disease incidence and mortality in Baja forests were lower. Although we quantitatively defined current Basin old‐growth forests – in terms of stand structure – we realize that our definition will differ from that of both past and future old‐growth forests unless management protocols are changed.     

Fire disturbance and forest structure in an old‐growth Pinus ponderosa forest,southern Cascades,USA

Questions: Did fire regimes in old‐growth Pinus ponderosa forest change with Euro‐American settlement compared to the pre‐settlement period? Do tree age structures exhibit a pattern of continuous regeneration or is regeneration episodic and related to fire disturbance or fire‐free periods? Are the forests compositionally stable? Do trees have a clumped spatial pattern and are clumps even‐ or mixed‐age? How might information from this old‐growth forest inform current restoration and management practices? Location: A 235‐ha old‐growth forest in the Ishi Wilderness, southern Cascade Mountains, California. Methods: Age, size, and spatial pattern of trees were quantified in seven stands. Fire history was reconstructed using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing fire history with age, size, and spatial structure of trees and identifying and measuring trees killed by two recent fires. Results: Species composition in plots was similar but density and basal area of tree populations varied. Age structure for P. ponderosa and Quercus kelloggii showed periods of episodic recruitment that varied among plots. Fire disturbance was frequent before 1905, with a median period between fires of 12 years. Fire frequency declined after 1905 but two recent fires (1990, 1994) killed 36% and 41% of mostly smaller diameter P. ponderosa and Q. kelloggii. Clusters of similar age trees occurred at scales of 28‐1018 m² but patches were not even‐aged. Interactions between tree regeneration and fire promoted development of uneven age groups of trees. Conclusions: Fire disturbance strongly influenced density, basal area, and spatial structure of tree populations. Fire exclusion over the last 100 years has caused compositional and structural changes. Two recent fires, however, thinned stands and created gaps favorable for Q. kelloggii and P. ponderosa regeneration. The effects of infrequent 20th century fire indicate that a low fire frequency can restore and sustain structural characteristics resembling those of the pre‐fire suppression period forest.     

Stand characteristics and biodiversity indicators along the productivity gradient in boreal forests: Defining a critical set of indicators for the monitoring of habitat nature quality

Abstract

We quantified the effects of anthropogenic disturbances on the structure and biodiversity of boreal forests on acidic soils and created a statistically supported rational set of indicators to monitor the stand “naturalness”. For that, we surveyed various traits of tree layer, understory, herb layer, forest floor and several widely accepted biodiversity epiphytic indicators in 252 old‐aged boreal stands in Estonia, mostly dominated by Scots pine or Norway spruce. Multifactorial general linear model analyses showed that many forest characteristics and potential indicators were confounded by the gradient of soil productivity (reflected by the forest site type), local biogeographic gradients and also by stand age. Considering confounding effects, boreal forests in a near‐natural state have more large‐diameter trees (diameter at breast height >40 cm) and larger variety of diameter classes, higher proportion of spruce or deciduous trees, a larger amount of coarse woody debris in various stages, a more closed tree canopy and denser understory than managed mature forests. By increasing light availability above the field layer, forest management indirectly increases the coverage of herbs and lichens on the forest floor but reduces the alpha‐ and beta‐diversity of herbs and the proportion of graminoids. Human disturbances reduce the relative incidence of many commonly accepted biodiversity indicators such as indicator lichens, woodpeckers, wood‐dwelling insects or fungi on trees. The test for the predictive power of characteristics reacting on disturbance revealed that only a fraction of them appeared to be included in a diagnostic easy‐to‐apply set of indicators to assess the nature quality of boreal forest: the amount of dead wood, the proportion of deciduous trees, the presence of specially shaped trees and woodpeckers and, as an indicator of disturbances, the forest herb Melampyrum pratensis. Many of these indicators have already been implemented in practice.     

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.     

Microsite variation and climate result in stand dynamics variation in pristine Pinus sylvestris mires: Evidence from a stand structure study

Questions: How do climate conditions and the site's ecohy‐ drological properties affect the age and size structure of natural Pinus sylvestris stands on pristine boreal mires? How do the long‐term stand dynamics on mires proceed as stands age? Do the mire stands reach a balanced, old‐growth stage? Location: Boreal mire forests in southern and northern Finland. Methods: Tree age and diameter distributions were analysed in 52 stands in two climate areas and in two mire site types with different ecohydrological properties. Temporal stand dynamics were examined by (1) comparing the graphs of the stands’ mean tree ages by diameter at breast height (1.3 m) classes and (2) describing the changes in stand characteristics and stand age and size structures as a function of stand dominant age in a chronosequence. Results: In the south, the DBH distributions were mostly unimodal and bell‐shaped in both site type groups. Age distributions were multimodal and flat in fully‐stocked sites but more uneven in sparsely forested composite sites. In the north, both the age and size distributions were clearly uneven in both site type groups. Tree age and size variation increased with stand age, but levelled out in the long term. Particularly in the south, the abundance of small trees decreased as stand age increased. Conclusions: The pine stands on pristine boreal mires are more dynamic than anticipated and are generally not characterised by a balanced, self‐perpetuating structure. Their dynamics reflect differences in climate and ecohydrology: on stocked sites in favourable boreal conditions, the stands showed structures typically resultant of inter‐tree competition processes that control tree growth and regeneration, whereas in harsh boreal climates, the tree regeneration process is ongoing diversifying the stand structure.     

Forests regenerating after clear-cutting function as habitat for bryophyte and lichen species of conservation concern

The majority of managed forests in Fennoscandia are younger than 70 years old but yet little is known about their potential to host rare and threatened species. In this study, we examined red-listed bryophytes and lichens in 19 young stands originating from clear-cutting (30-70 years old) in the boreal region, finding 19 red-listed species (six bryophytes and 13 lichens). We used adjoining old stands, which most likely never had been clear-cut, as reference. The old stands contained significantly more species, but when taking the amount of biological legacies (i.e., remaining deciduous trees and dead wood) from the previous forest generation into account, bryophyte species number did not differ between old and young stands, and lichen number was even higher in young stands. No dispersal effect could be detected from the old to the young stands. The amount of wetlands in the surroundings was important for bryophytes, as was the area of old forest for both lichens and bryophytes. A cardinal position of young stands to the north of old stands was beneficial to red-listed bryophytes as well as lichens. We conclude that young forest plantations may function as habitat for red-listed species, but that this depends on presence of structures from the previous forest generation, and also on qualities in the surrounding landscape. Nevertheless, at repeated clear-cuttings, a successive decrease in species populations in young production stands is likely, due to increased fragmentation and reduced substrate amounts. Retention of dead wood and deciduous trees might be efficient conservation measures. Although priority needs to be given to preservation of remnant old-growth forests, we argue that young forests rich in biological legacies and located in landscapes with high amounts of old forests may have a conservation value.     

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.     

Dead Wood is Buried and Preserved in a Labrador Boreal Forest

Large amounts (36.4 Mg ha⁻¹ or 179 m³ ha⁻¹) of buried dead wood were found in overmature (146–204-year-old) black spruce (Picea mariana (Mill.) B.S.P.) forests in the high boreal region of eastern Canada. Amounts of this size indicate that burial reduces rates of wood decay producing an important component of long-term carbon (C) storage. Radiocarbon-derived ages of black spruce stems buried near the bottom of the organic soil horizon at three old-growth sites were up to 515 years old. Together with information on current stand age, this suggests that the stems have been dead for more than 250 years. Most aboveground dead wood decays or becomes fragmented within about 70 years of tree death in these forests. The presence of old yet well-preserved buried wood suggests that decay rates are greatly reduced when downed dead wood is quickly overgrown by moss. Thus, the nature and type of ground-layer vegetation influences the accumulation of organic matter in these forests. This process of dead wood burial and the resultant addition to a large and long-enduring belowground C pool should be considered when estimating dead wood abundance for habitat or forest C accounting and cycling.     

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.     

Succession in sub‐boreal forests of West‐Central British Columbia

Abstract. Structural and compositional changes were analysed over the course of 400+ yr of post‐fire succession in the sub‐boreal forests of west‐central British Columbia. Using a chronosequence of 57 stands ranging from 11 to 438 yr in age, we examined changes in forest structure and composition with complementary PCA and DCA ordination techniques. To determine stand ages and timing of tree recruitment, approximately 1800 trees were aged. Most early successional forests were dominated by Pinus contorta, which established rapidly following fire. Abies lasiocarpa and Picea glauca × engel‐mannii were also able to establish quickly, but continued to establish throughout the sere. Few Pinus contorta survived beyond 200 yr, resulting in major changes in forest structure. In some stands P. contorta never established, which led to considerable variation among stands less than 200 yr old. The oldest forests converged on dominance by Abies lasiocarpa. Vascular plant diversity decreased during succession whereas canopy structure became more complex as gap dynamics developed. Although these sub‐boreal forests contain few tree species, successional changes were pronounced, with structure changing more than composition across the chronosequence.     

Structural characteristics and aboveground biomass of old‐growth spruce–fir stands in the eastern Carpathian mountains,Ukraine

Abstract

Temperate old‐growth forests are known to have ecological characteristics distinct from younger forests, but these have been poorly described for the remaining old‐growth Picea abies–Abies alba forests in the eastern Carpathian mountains. In addition, recent studies suggest that old‐growth forests may be more significant carbon sinks than previously recognized. This has stimulated interest in quantifying aboveground carbon stocks in primary forest systems. We investigated the structural attributes and aboveground biomass in two remnant old‐growth spruce–fir stands and compared these against a primary (never logged) mature reference stand. Our sites were located in the Gorgany Nature Reserve in western Ukraine. Overstory data were collected using variable radius plots; coarse woody debris was sampled along line intercept transects. Differences among sites were assessed using non‐parametric statistical analyses. Goodness‐of‐fit tests were used to evaluate the form of diameter distributions. The results strongly supported the hypothesis that old‐growth temperate spruce–fir forests have greater structural complexity compared to mature forests, including higher densities of large trees, more complex horizontal structure, and elevated aboveground biomass. The late‐successional sites we sampled exhibited rotated sigmoid diameter distributions; these may reflect natural disturbance dynamics. Old‐growth Carpathian spruce–fir forests store on average approximately 155–165 Mg ha^?1 of carbon in aboveground tree parts alone. This is approximately 50% higher than mature stands. Given the scarcity of primary spruce–fir forests in the Carpathian region, remaining stands have high conservation value, both as habitat for late‐successional species and as carbon storage reservoirs.     

Role of the Siberian flying squirrel as an umbrella species for biodiversity in northern boreal forests

One of the potentially useful indirect shortcut methods in biodiversity conservation is the umbrella species concept. An umbrella species can be seen relatively demanding for the size of the area and probably also for certain habitat types: conservation management for the umbrella species would thus encompass other species preferring similar habitats but with smaller area requirements. As such, it has a comprehensive spatial aspect for landscape planning. We tested the role of the Siberian flying squirrel (Pteromys volans) as an umbrella species for wood dependent species among red-listed and old-growth forest associated polypores, epiphytic lichens and beetles. Flying squirrels inhabit home ranges of several to tens of hectares, and prefer mature spruce-dominated (Picea abies) mixed forests, which often have high amounts of dead wood. We carried out species surveys and trappings during 1 year from 20 mature spruce-dominated forest stands (altogether 162 ha), of which 12 were occupied by the flying squirrel. The amount of dead wood was higher in occupied stands than in unoccupied stands. We also found a tendency for a higher number of species and number of records in occupied stands, a relationship mostly due to the polypore species. The presence of the flying squirrel may reflect the habitat availability for species depending on dead and living wood, and assist in site selection of conservation areas. We suggest that the flying squirrel has potential as an umbrella species to partly enhance maintenance of biodiversity in northern boreal forests in Finland.     

Conservation management of Pinus palustris ecosystems from a landscape perspective

Question: Do case studies from silvicultural and restoration studies and applied conservation management in second‐growth Pinus palustris stands provide unique insights for conservation models? Methods: A review of management paradigms that conserve the high biological diversity and rare species, drawn from characteristics in both second‐growth and old‐growth stands, is presented for fire‐maintained Pinus palustris (longleaf pine) forests. Results: A common assumption that old‐growth stands provide the primary information for the development of conservation management strategies de‐emphasizes lessons learned from second‐growth and restoration studies. Primary conservation management goals for the Pinus palustris ecosystem include the perpetual regeneration of the fire‐maintained forest and conservation of the characteristically high biological diversity and rare species. Several attributes, such as a sustained population of Picoides borealis (red‐cockaded woodpecker), Aristida stricta (wiregrass)‐dominated ground cover, and undisturbed upland‐wetland ecotones, can predict a diverse and ecologically functional ecosystem. Such indicators are linked to critical structural and functional features of the system and reflect previous land management histories that suggest sustainable approaches. Conclusions: A traditional definition of ‘old‐growth’ relying on overstorey may be limited in describing important features of healthy, diverse Pinus palustris ecosystems. Some characteristics are significantly more important for maintenance of diversity than age of the trees and these features may be present in old‐ or second‐growth forests. We advocate that the management history, structural characteristics and landscape context of stands that harbour desirable conservation attributes (red‐cockaded woodpeckers, wiregrass, gopher tortoises and undisturbed upland‐wetland ecotones) can be used as indicators to identify important conservation and forest management principles.     

Diversity of saproxylic bryophytes in old‐growth and managed beech forests in the central Balkans

Abstract

The diversity of saproxylic bryophyte species in beech forest stands from the wide region of the central Balkans (i.e. Serbia and Montenegro) was studied, and this study is the first of such a type in SE Europe. Comparison of preserved old‐growth and managed forests were made. Bryophyte species diversity is higher in primeval forest stands where the spectra of dead wood in various decaying stages of its dynamics are present. The ecological group of epixylic specialists is predominant among the bryophytes recorded. Threatened bryophyte species occur in old‐growth beech stands. The dead wood as habitat together with some other factors are extremely important for the surviving of epixylic bryophyte; so these species can be used as bioindicator bryophyte species of old‐growth or managed and structured forest ecosystems.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Natural and anthropogenic influences on coarse woody debris stocks in Nothofagus–Araucaria forests of northern Patagonia,Argentina

Coarse woody debris (CWD) is an important element driving ecological processes, strengthening ecosystem resilience and for biodiversity within forest ecosystems. However, the abundance and distribution of CWD and their relation to natural and human factors are poorly known in southern South America. In this work we studied the density and volume of CWD types in Nothofagus–Araucaria stands in northern Patagonia (Neuquén – Argentina) and relationships with forest composition and structure. We also studied their relationships with fire history, topography and human‐related variables. Twenty‐three stands with Nothofagus pumilio, Nothofagus antarctica and/or Araucaria araucana were sampled to estimate quantities of logs, snags and dead branches using the planar‐intersect method. CWD density and volume in these forests were moderate and varied across the landscape with a spatial pattern determined by biotic, abiotic and human use–related variables. Mean CWD volume was 52.9 m³ ha^?1 (range: 1.6–143.7) and significantly varied among forest types and watersheds. CWD was positively related to dbh, tree height and slope, but negatively related to tree density. CWD was clearly influenced by composition and structural characteristics of stands, where the tree species traits had an important role. As well, the observed amount and type of CWD, whereby most of the stands showed low levels of old (pre‐disturbance) logs/snags and poor new inputs of deadwood, may be explained by fire frequency. Firewood gathering and livestock grazing negatively affected deadwood stocks and topography counteracts this effect by limiting human access. Fire disturbance history, windthrow and dieback pulses produced by insect outbreaks and human access seemed to be the main causes that best explained CWD spatial distribution and abundance patterns in north‐western Patagonian forests.     

The Role of Wood Ants (Formica rufa group) in Carbon and Nutrient Dynamics of a Boreal Norway Spruce Forest Ecosystem

Wood ants (Formica rufa group) are regarded as keystone species in boreal and mountain forests of Europe and Asia by their effect on ecosystem carbon (C) and nutrient pools and fluxes. To quantify the impact of their activity on boreal forest ecosystems, C, nitrogen (N), phosphorus (P), potassium (K) and calcium (Ca) pools and fluxes in wood ant nests (WAN), and soil were assessed along a 5-, 30-, 60-, and 100-year-old Norway spruce (Picea abies L. Karsten) dominated successional gradient in eastern Finland. Amounts of C and nutrients in WAN increased with stand age, but contained less than 1% of total C and nutrient pools in these stands. The CO₂-efflux from nests was also insignificant, as compared to CO₂-efflux from the forest floor. Annually, the amount of C brought by wood ants into their nests as honeydew, prey and nest-building materials ranged from 2.7 to 49.3 kg ha^?1 C, but this is only 0.1–0.7% of the combined net primary production of trees and understorey in boreal forests. The difference between wood ant nest C inputs and outputs was very small in the younger-aged stands, and increased in the older stands. Carbon accumulation rates in nests over a 100 year period are estimated to be less than 10 kg ha^?1 a^?1. In contrast to C, annual inputs of N, P, and K are larger compared to wood ant nest nutrient pool size, ranging from 3 to 6% of the annual tree stand and understorey uptake. This indicates a more rapid turnover and transport of N, P, and K out of WAN, and suggests that wood ants increase the cycling rate of these nutrients in boreal forests.