Comparisons of arthropod and avian assemblages in insecticide-treated and untreated eastern hemlock (Tsuga canadensis [L.] Carr) stands in Great Smoky Mountains National Park,USA

Great Smoky Mountains National Park is using systemic imidacloprid to treat eastern hemlocks (Tsuga canadensis [L.] Carr.) infested with the exotic insect, hemlock woolly adelgid (Adelges tsugae Annand). This study investigated effects of these treatments on insectivorous birds and hemlock canopy arthropod assemblages in the context of food availability for insectivorous birds. Six pairs of treated and untreated hemlock sites were studied in 2007. Territories of three hemlock-associated Neotropical migratory foliage-gleaning bird species were mapped in these six sites, and relationships between bird territory density and hemlock foliar density were examined. Canopy arthropods were sampled by clipping mid-canopy hemlock branches in each paired site. Arthropods were identified to order or suborder and categorized into bird prey guilds and non-target herbivorous insect guilds. Despite being treated within the previous two years, there were no differences in hemlock woolly adelgid infestation between treated and untreated sites. This may reflect recovery or because the imidacloprid is slow-acting. Bird densities also did not differ between treated and untreated sites but were positively related to branch foliage mass, implying a preference in these birds for well-foliated hemlocks. A total of 10,219 hemlock woolly adelgids, and a total of 906 other arthropods from 16 orders were collected. There were no differences in species richness, abundance, or species composition between treated and untreated sites for total arthropods, or for immature arthropods ≥ 3 mm. In contrast, non-target herbivorous Hemiptera and larval Lepidoptera were significantly reduced in treated hemlocks. Although larval Lepidoptera are primary prey for insectivorous foliage-gleaning birds, the similarity in bird densities between treated and untreated sites suggests the birds are able to find other food resources in the mixed hemlock-deciduous stands where the study sites were located. Therefore, controlling hemlock woolly adelgid-induced defoliation through use of imidacloprid may have short-term benefits for hemlock-associated birds. While imidacloprid treatments did not appear to be currently affecting most arthropods, primary prey guilds should be monitored for long-term declines that could impact hemlock-associated birds.     

Changes in light levels and stream temperatures with loss of eastern hemlock (Tsuga canadensis) at a southern Appalachian stream: Implications for brook trout

The exotic invasive insect, hemlock woolly adelgid (Adelges tsugae Annand), is causing mortality in eastern hemlocks (Tsuga canadensis [L.] Carr.) throughout the eastern U.S. Because hemlocks produce dense shade, and are being replaced by hardwood species that produce less shade, their loss may increase understory light levels. In the southern Appalachians, increases in light could increase stream temperatures, threatening species such as brook trout (Salvelinus fontinalis). We studied changes in light and stream temperature with eastern hemlock decline at a headwater southern Appalachian brook trout stream. Our results indicate that stream light levels have increased significantly with adelgid infestation. Leaf-on light levels are currently significantly higher (P < 0.02) in plots containing high basal areas of hemlock (mean global site factor (GSF)(SE) = 0.267(0.01)) compared with plots containing no hemlock (mean GSF(SE) = 0.261(0.01)), suggesting that increases in light have occurred with hemlock decline. The Normalized Difference Vegetation Index (NDVI), a remotely sensed metric of vegetation density, decreased with hemlock decline from 2001 to 2008. In 2001, NDVI showed no relationship (R² = 0.003; F = 0.14; P = 0.71) with hemlock basal area, but by 2008, there was a significant negative relationship (R² = 0.352; F = 19.55; P < 0.001) between NDVI and hemlock basal area. A gap experiment showed that light levels may increase by up to 64.7% more (mean increase in GSF = 27.5%) as hemlocks fall, creating gaps in the canopy. However, stream temperatures did not increase with hemlock decline during the study period, and we found that ground water inputs have a stronger influence on water temperature than light levels at this site. Linear regression showed a significant negative relationship between water temperature and proximity to ground water sources (R² = 0.451; F = 13.14; P = 0.002), but no relationship between water temperature and light levels (R² < 0.02; P > 0.05). In addition, by comparing light levels between plots containing hemlock and those containing only hardwoods, we found that if hemlocks are replaced by hardwoods, light levels under an all-hardwood canopy (mean GSF(SE) = 0.240(0.005)) are unlikely to be higher than they are under the current forest (mean GSF(SE) = 0.254(0.007)). These results suggest that loss of hemlock along southern Appalachian headwater streams could have short-term impacts on light levels, but that long-term changes in light levels, increases in water temperature, and adverse effects on brook trout may be unlikely.     

Water use and carbon exchange of red oak- and eastern hemlock-dominated forests in the northeastern USA: implications for ecosystem-level effects of hemlock woolly adelgid

Water use and carbon exchange of a red oak-dominated (Quercus rubra L.) forest and an eastern hemlock-dominated (Tsuga canadensis L.) forest, each located within the Harvard Forest in north-central Massachusetts, were measured for 2 years by the eddy flux method. Water use by the red oak forest reached 4 mm day(-1), compared to a maximum of 2 mm day(-1) by the eastern hemlock forest. Maximal carbon (C) uptake rate was also higher in the red oak forest than in the eastern hemlock forest (about 25 versus 15 micromol m(-2) s(-1)). Sap flux measurements indicated that transpiration of red oak, and also of black birch (Betula lenta L.), which frequently replaces eastern hemlock killed by hemlock woolly adelgid (Adelges tsugae Annand.), were almost twice that of eastern hemlock. Despite the difference between species in maximum summertime C assimilation rate, annual C storage of the eastern hemlock forest almost equaled that of the red oak forest because of net C uptake by eastern hemlock during unusually warm fall and spring weather, and a near-zero C balance during the winter. Thus, the effect on C storage of replacing eastern hemlock forest with a forest dominated by deciduous species is unclear. Carbon storage by eastern hemlock forests during fall, winter and spring is likely to increase in the event of climate warming, although this may be offset by C loss during hotter summers. Our results indicate that, although forest water use will decrease immediately following eastern hemlock mortality due to the hemlock woolly adelgid, the replacement of eastern hemlock by deciduous species such as red oak will likely increase summertime water use over current rates in areas where hemlock is a major forest species.     

Forest management conditioning ground ant community structure and composition in temperate conifer forests in the Pyrenees Mountains

The search for indicators to monitor management impact on biodiversity is a crucial question because management practices promote changes in community structure and composition of different animal groups. This study explores the effect of widely conducted management practices (forest logging and livestock) in Pinus uncinata forests in the Pyrenees range (NE Spain) on the structure and composition of ground ant communities compared to those of old-growth stands. Forest structure clearly differed in stands with different forest managements. These stands managed for different uses also showed marked differences in structure and composition of ground ant communities. There was a great dominance of a single species, Formica lugubris, which accounted for 99% of ants collected in pitfall traps. Rarefaction curves indicated that species richness was highest in old-growth stands and lowest in even-aged ones, with woodland pasture stands showing an intermediate value. Classification methods allowed us to identify two groups of species: six species related to old-growth plots and three species (including F. lugubris) associated to managed stands. Habitat structure played an important role in determining the structure of ant communities: forests with high tree density but low basal area were the most favourable forest type for F. lugubris, while the abundance of the remaining ant species was negatively affected by the abundance of F. lugubris and by tree cover.     

Optimizing the joint production of timber and bilberries

Berries and mushrooms are increasingly appreciated products of Finnish forests. Therefore, there is a need to integrate them in silvicultural planning. Bilberry (Vaccinium myrtillus L.) is an economically important wild berry that is widely collected for household consumption and sale in North Karelia, Finland. In this study, bilberry yield models developed recently were included in a stand growth simulator and the joint production of timber and bilberry was optimized by maximizing soil expectation value (SEV) with 3% discounting rate, assuming that 75% of the bilberry yield is harvested. The effect of bilberry production on the optimal stand management increased with increasing bilberry price. With high bilberry prices (4–8 € kg⁻¹) it was optimal to manage the mixed stand of Scots pine, Norway spruce and birch, and the pure stand of Norway spruce so as to promote bilberry production. In the Scots pine stand, where bilberry yields are higher, bilberry production affected optimal stand management already with a price of 2 € kg⁻¹. Compared to timber production, joint production led to longer rotation lengths, higher thinning intensities, more frequent thinnings, and higher share of Scots pine in the mixed stand. The contribution of bilberries to the total SEV increased with increasing bilberry price and discounting rate. In the mixed stand and pine stand the SEV of bilberry production, calculated with 3% discounting rate, exceeded the SEV of timber production when bilberry price was 4 € kg⁻¹.With 4% discounting rate this happened already with bilberry price of 2 € kg⁻¹. It was concluded that forest management which promotes bilberry yields is the most profitable in pine stands where the potential bilberry yields are high.     

Response of bark beetles and their natural enemies to fire and fire surrogate treatments in mixed-conifer forests in western Montana

Four treatments (control, burn-only, thin-only, and thin-and-burn) were evaluated for their effects on bark beetle-caused mortality in both the short-term (one to four years) and the long-term (seven years) in mixed-conifer forests in western Montana, USA. In addition to assessing bark beetle responses to these treatments, we also measured natural enemy landing rates and resin flow of ponderosa pine (Pinus ponderosa) the season fire treatments were implemented. All bark beetles were present at low population levels (non-outbreak) for the duration of the study. Post-treatment mortality of trees due to bark beetles was lowest in the thin-only and control units and highest in the units receiving burns. Three tree-killing bark beetle species responded positively to fire treatments: Douglas-fir beetle (Dendroctonus pseudotsugae), pine engraver (Ips pini), and western pine beetle (Dendroctonus brevicomis). Red turpentine beetle (Dendroctonus valens) responded positively to fire treatments, but never caused mortality. Three fire damage variables tested (height of crown scorch, percent circumference of the tree bole scorched, or degree of ground char) were significant factors in predicting beetle attack on trees. Douglas-fir beetle and pine engraver responded rapidly to increased availability of resources (fire-damaged trees); however, successful attacks dropped rapidly once these resources were depleted. Movement to green trees by pine engraver was not observed in plots receiving fire treatments, or in thinned plots where slash supported substantial reproduction by this beetle. The fourth tree-killing beetle present at the site, the mountain pine beetle, did not exhibit responses to any treatment. Natural enemies generally arrived at trees the same time as host bark beetles. However, the landing rates of only one, Medetera spp., was affected by treatment. This predator responded positively to thinning treatments. This insect was present in very high numbers indicating a regulatory effect on beetles, at least in the short-term, in thinned stands. Resin flow decreased from June to August. However, resin flow was significantly higher in trees in August than in June in fire treatments. Increased flow in burned trees later in the season did not affect beetle attack success. Overall, responses by beetles to treatments were short-term and limited to fire-damaged trees. Expansions into green trees did not occur. This lack of spread was likely due to a combination of high tree vigor in residual stands and low background populations of bark beetles.     

Impacts of hemlock looper defoliation on growth and survival of balsam fir, black spruce and white birch in Newfoundland, Canada

Hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) is an economically damaging defoliator that undergoes periodic outbreaks in Newfoundland, Canada. It defoliates and causes extensive tree mortality to its primary host, balsam fir (Abies balsamea [L.] Mill.). We quantified tree survival using data from permanent sample plots (PSPs) and growth reduction or release using dendrochronology, and related these impacts to defoliation severity determined from annual aerial defoliation survey data. Such impact relationships are necessary as a fundamental input to a Decision Support System. Growth and survival of balsam fir, black spruce (Picea mariana (Mill.) B.S.P.) and white birch (Betula papyrifera Marsh.) were assessed from 1996 to 2008 in 48 Newfoundland Forest Service PSPs, selected based on four classes of defoliation severity. Two years of severe (71-100%) defoliation resulted in almost complete mortality for balsam fir, 10 years after defoliation, whereas survival was 70-80% for black spruce and white birch. Lower defoliation severity (1-2 years of moderate (31-70%) or 1 year of severe) resulted in approximately 60% survival for balsam fir and no reduction in survival for black spruce and white birch. Maximum growth reduction of balsam fir was 10-15% with 1 year of moderate-severe defoliation, 35-40% with 2 years of moderate defoliation, and about 50% with 2 years of severe defoliation. Growth recovered to pre-defoliation rates 5 years after defoliation ceased in all severity classes. Growth reduction and recovery of black spruce were more variable and lower than for balsam fir, and white birch exhibited only minor (<10%) growth reduction during the defoliation year or 1 year after defoliation. Control measures should focus on avoiding severe defoliation for two consecutive years.     

Snag longevity of Douglas-fir,western hemlock,and western redcedar from permanent sample plots in coastal British Columbia

Snags are important both as structural components and as animal habitat in forests, but abundance is often low and their dynamics poorly understood in young, managed stands. Using a large data set of 19,622 snags from permanent plots in second-growth forests of coastal British Columbia, we modeled snag longevity (time from tree mortality to snag fall) for three species: Douglas-fir (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla), and western redcedar (Thuja plicata). Snag longevity was strongly related to species and snag size (diameter): the median snag longevity was 16 years for Douglas-fir, 11 years for hemlock and 5 years for redcedar. Western redcedar was predominantly in the subcanopy and its rapid fall rate was related to the small size of its snags. In addition to diameter, other attributes (height to diameter ratio, height, and live crown ratio before death) contributed significantly to models for one or two of the species. However, site level variables did not contribute significantly to any of the models. Snags greater than 50 cm diameter, especially Douglas-fir snags, have the potential for persistence well beyond 20 years in these second-growth forests, and could be important for wildlife.     

Effects of non-native spruce plantations on small mammal communities in subarctic birch forests

Voles and shrews are key species in northern forest ecosystems. Thus, it is important to quantify to what extent new forestry practices such as planting of non-native tree species impact these small mammals. In northern Norway stands of coastal subarctic birch forests have increasingly been converted to non-native spruce stands during the last century. This leads to changes in the forest floor vegetation and soil conditions that can be expected to negatively impact the community of ground-dwelling small mammals. In this 10-year trapping study we contrasted seasonal small mammal population abundances in spruce plantations with four birch forest varieties. Six different small mammal species were trapped (in descending order of abundance; common shrew Sorex araneus, red vole Myodes rutilus, field vole Microtus agrestis, grey-sided vole M. rufocanus, pygmy shrew S. minutus and water shrew Neomys fodiens). None of the voles appeared to exhibit temporal dynamics resembling population cycles. The three most numerous species were clearly less abundant in the spruce plantations compared to the other forest types. Autumn abundances were most impacted by spruce plantations, indicating that growth rates in the reproductive season were more influenced than winter declines. Species associated with productive forest habitats (i.e. field vole and common shrew) were most impacted by tree species conversion. Still young spruce plantations inter-mixed with birch trees and the ecotone habitat, sustained small mammal abundances comparable to the native birch forests. This implies that managing spruce plantations to maintain a mix of different tree species and high spatial heterogeneity (i.e. more ecotones), will reduce the negative impacts on the small mammal community. On the contrary, if young spruce plantations, as they age become spruce monocultures covering larger parts of the landscapes than they do presently, the negative effects on small mammal communities may be larger than observed in the present study.     

Climate and historical stand dynamics in the tropical pine forests of northern Thailand

Forest recruitment is the outcome of local- and regional-scale factors such as disturbances and climate. The relative importance of local- and regional-scale factors will determine the spatial scale at which temporal pulses of recruitment occur. In seasonal tropical forests, where the annual dry-season is a critical bottleneck to seedling survival, multi-year periods of relatively cool, wet dry seasons may be required for successful tree recruitment. Consequently, when such conditions are present, region-wide synchronisation of recruitment may occur. To examine the case for regional synchronisation of forest dynamics in the seasonal tropical pine forests of northern Thailand, we investigated forest age structures at three spatial scales: stand, site and region. We compared forest age structures with instrumental climatic records beginning in 1902. We found significant statistical evidence of synchronous recruitment at the stand- and site-scales, but not at the regional-scale. While correlations between recruitment and climate were not statistically significant, recruitment success was often linked to favourable climatic conditions. For example, recruitment at all sites was associated with multi-year periods of cool-wet dry seasons. The lack of significant correlations between recruitment and climate appears to reflect complex interactions among local disturbance history, regional climate variability and pine recruitment.     

Restoration of dry tropical forests in Central America: A review of pattern and process

Much information on restoration and management exists for wet tropical forests of Central America but comparatively little work has been done in the dry forests of this region. Such information is critical for reforestation efforts that are now occurring throughout Central America. This paper describes processes of degradation due to land use and provides a conceptual framework for the restoration of dry tropical forest. Most of this forest type was initially harvested for timber and then cleared for cattle in the last century (1930-1970). Only 1.7% remains largely restricted to infertile soils and remote areas on the Pacific coastal side of Panama, Costa Rica, Nicaragua and Mexico. These cleared areas are again in a state of transition due to a combination of decreasing land productivity, and land speculation for tourism development. Some farms have been sold to new landowners who are interested in reforesting to increase biodiversity and forest cover. Attempts have therefore been made to reforest by protecting the land from fire and cattle, by supplementing natural regrowth with enrichment planting, or through use of tree plantations. Experimental studies have demonstrated the ability of these lands to grow back to forests because of native species ability to sprout after cutting, and the capacity of remnant trees in field and riparian zones to provide seeds and to moderate edge environment for seed germination and seedling establishment. However, research also shows that on sites with long histories of land clearance, species diversity will remain low with functional groups missing unless some active management occurs. Under-planting with late-successional native tree species can add structure and diversity; enrichment planting with large-fruited shade-intolerant species can initiate new islands of more diverse regeneration beneath their canopies; and plantings of fast-growing, nitrogen-fixing trees that provide light canopy shade can moderate the environment below, promoting regeneration establishment of late-successional species. Plantations are the only option for lands that have lost almost all remnants of native forest, and where soils and vegetation have changed to new states of structure and function. Conversion of pastures to tree plantations that can facilitate natural regeneration beneath them is appropriate when pastures are prone to fire and/or lack immediate seed sources nearby. After the grasses have been shaded out, natural recruitment can slowly occur over a 10-15 years period. Under-planting of shade-tolerant late-successional species can supplement species composition and structure.     

Persistent effects of fire severity on early successional forests in interior Alaska

There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

The lateral spread of tree root systems in boreal forests: Estimates based on N uptake and distribution of sporocarps of ectomycorrhizal fungi

In nutrient poor environments, such as boreal forests, many of the most important interactions between plants take place belowground. Here, we report the results of two approaches to obtain estimates of the lateral spread of tree roots.     

Sap flow measurements reveal influence of temperature and stand structure on water use of Eucalyptus regnans forests

We examined water use by maturing Eucalyptus regnans, growing with or without an mid-storey stratum of Acacia spp. (Acacia dealbata or A. melanoxylon), for >180 consecutive days. Study sites were located in the Upper Yarra catchment area in south-eastern Australia. Depending on their contribution to stand basal area, mid-storey Acacia spp. increased total stand water use by up to 30%. Monthly water use in such stands reached more than 640,000 L ha⁻¹ (compared to 545,000 L ha⁻¹ in stands where acacias were absent) in early spring. Water use was curvilinearly related to sapwood area of Acacia spp. and logistically related to sapwood area of E. regnans. Water use of all three species showed a strong relation to daily maximum air temperatures. Distinct and simple relationships provide clear guides to the likely impacts of climate change and forest management on water yield. We compared a traditional up-scaling approach, from individual tree water use to stand water use, to a new approach that incorporates variation in temperature. Development of this approach can lead to greater precision of stand water use estimates – and in turn catchment water yield – under current climate change scenarios, which predict a rise in air temperatures of 0.6–2.5 °C by 2050 for the study area. Our temperature-dependent approach suggests that under conditions of non-limiting water availability, stand water use will rise by 2% for every 0.25 °C increase in maximum air temperatures during winter, and possibly more than that during summer.     

Driving factors for natural tree rejuvenation in anthropogenic pine (Pinus sylvestris L.) forests of NE Germany

The rejuvenation ecology of three main tree species in anthropogenic pine (Pinus sylvestris L.) forests is explored in our study. We focus on the scale of micro-plots, which provide the safe sites for tree rejuvenation. We thrive on the multi-factorial relationship of tree establishment and driving ecological factors using a large dataset from pine stands in NE Germany and applying multivariate analyses. The success of the establishment of the investigated focal tree species Fagus sylvatica L., Quercus petraea Liebl. and Pinus sylvestris L. is, on general, mostly affected by three factors, i.e. water balance of the upper soil layers, browsing pressure, and diaspore sources. Our investigations on the micro-plot scale revealed species-specific differences. For beech saplings <50 cm growth height, primarily the availability of water, indicated by available water capacity (AWC), thickness, quality, and structure of the organic layer, silt and humus content in the topsoil, and the lack of a dense competitive herb layer, were identified as most important factors. On the contrary, oak seems hardly be restricted by hydrologic and/or trophic deficits in the topsoil or humus layer. In conclusion and comparison to Fagus sylvatica L., we assume for Quercus petraea Liebl. advantages in natural regeneration processes under sub-continental climate conditions and thus under the scenarios of climate change. Pinus sylvestris L. regeneration in our investigation area occurs only in a narrow niche. We conclude with regard to future forest development and the objective of stand conversion with low management intensity that oak should be favoured within natural stand regeneration.     

Burning and mowing as habitat management for capercaillie Tetrao urogallus: An experimental test

In Scots pine Pinus sylvestris forests, the important ecological effects of natural fires could be emulated using prescribed fire. Species that may benefit from fire effects include capercaillie Tetrao urogallus, a large forest grouse. A key component of forest habitats for capercaillie is the ericaceous shrub, bilberry Vaccinium myrtillus, which is eaten by capercaillie, and supports abundant arthropods, taken by young chicks. We carried out an experiment testing whether prescribed burning would be a valuable technique for capercaillie habitat management. The study took place at Abernethy Forest, the largest ancient native pinewood in Britain, and a key capercaillie site, holding c 8-20% of the British population. Prescribed fire in woodland is highly novel in Britain. We therefore also tested mowing, which might replicate some fire effects more cheaply and safely. Twenty-five experimental blocks were established within open pine stands with ground vegetation including bilberry, but dominated by heather Calluna vulgaris. Each block held three 700 m² plots, randomly assigned to control, mow and burn. Vegetation, arthropods and capercaillie dung were monitored over a 7-year period, including 1 year prior to treatment. Mean bilberry cover, initially around 12%, increased in mown and burnt areas, but there were also increases in controls, following unusual natural die-back of heather. By the sixth season after treatment, bilberry cover was significantly higher in burnt and mown areas than controls, averaging 27% (95% confidence intervals 24-30), compared to 20% (19-21) in controls. Biomass of spiders, an important dietary group for capercaillie chicks, as measured by pitfall trapping, was significantly higher in burnt and mown plots than controls, by about 56% (38-76). However, biomass of caterpillars, often considered a more important dietary group, did not show clear differences between treatments. An alternative analysis was used to ‘statistically remove’ natural heather die-back; this enhanced the treatment differences in bilberry cover and spider biomass. Capercaillie dung counts suggested that burnt, and especially mown areas, had more summer capercaillie usage than controls. Capercaillie conservation at sites similar to Abernethy is likely to benefit from either prescribed fire or mowing, because these techniques increase bilberry and spider abundance. This study illustrates the value of collaboration between researchers and land-managers, in developing and testing novel management techniques. We support the idea that ‘dominance reduction’, delivered through managed disturbance, offers a general principle to guide land-managers wishing to maintain biodiversity, particularly where key species, like capercaillie, are strongly associated with sub-dominant plant species like bilberry.     

Epiphyte diversity and biomass loads of canopy emergent trees in Chilean temperate rain forests: A neglected functional component

We document for the first time the epiphytic composition and biomass of canopy emergent trees from temperate, old-growth coastal rainforests of Chile (42°30′S). Through tree-climbing techniques, we accessed the crown of two large (c. 1 m trunk diameter, 25–30 m tall) individuals of Eucryphia cordifolia (Cunoniaceae) and one large Aextoxicon punctatum (Aextoxicaceae) to sample all epiphytes from the base to the treetop. Epiphytes, with the exception of the hemi-epiphytic tree Raukaua laetevirens (Araliaceae), were removed, weighed and subsamples dried to estimate total dry mass. We recorded 22 species of vascular epiphytes, and 22 genera of cryptogams, with at least 30 species of bryophytes, liverworts and lichens. The dominant vascular epiphytes were Fascicularia bicolor (Bromeliaceae), Raukaua laetevirens, Sarmienta repens (Gesneriaceae), and filmy ferns (Hymenophyllaceae). Epiphyte loads per tree ranged between 134 and 144 kg dry mass, with 60–70% water. The hemi-epiphytic tree R. laetevirens added between 1 and 2.6 t of dry mass to each host tree. A main component of epiphyte biomass, making 70% of the weight, was detritus and roots, while leaves, stems, and fronds made up the remaining 30%. Emergent trees hold a high proportion of the regional diversity of epiphytes: 33% of all flowering epiphytes, and 50% of all filmy ferns described for Chilean temperate forests. Dry epiphyte biomass associated only with the emergent E. cordifolia trees in coastal forests was estimated in 10 t/ha. Epiphyte biomass may store up to 300 l of water in each emergent tree, and add 40–150% of photosynthetic biomass to the tree crowns. Based on this evidence, epiphytes may play key but generally neglected roles in ecosystem carbon uptake, water storage, and nutrient cycling. Moreover, emergent trees represent nuclei of biodiversity and ecosystem functions distributed throughout mature forests. Forest management should recognize large trees as significant management units for the preservation of biodiversity and ecological functions.     

The impact of black cottonwood on soil fertility in coastal western hemlock forest

Black cottonwood (Populus trichocarpa Torr. and Gray) is a deciduous tree species that extends from Alaska through coastal regions of western Canada into the northwestern United States and as far south as Baja California. We examined the influence of black cottonwood on soil fertility within a forest dominated by Douglas-fir [Pseudotsuga menziessi (Mirb.) Franco], western hemlock [Tsuga heterophylla (Raf.) Sarg], and western red cedar (Thuja plicata Donn ex. D. Don.). Six circular 0.008 ha plots with a single cottonwood tree in the center of conifers were paired with six conifer plots (of the same size) without cottonwood. Litterfall, litter decomposition, properties of forest floor and mineral soil, and N mineralization were compared between plot types. Cottonwood litter had higher concentrations of almost all elements relative to conifer litter. Mass loss did not differ between cottonwood and fir/hemlock litter on cottonwood sites. Twice the amount of mull-like humus form (vermimull and mullmoder, 56%) was found in cottonwood plots compared to 28% in conifer plots. Higher pH (4.4) was found in the forest floor under cottonwood compared to conifer (3.9). Total N concentration (3.33 g/kg) and base saturation (68%) were higher in the mineral soil under cottonwood compared to conifers (2.98 g/kg total N and 50% base saturation). Net ammonification and net mineralization were both lower under cottonwood. These results suggest a variable effect of cottonwood on soil fertility within coastal western hemlock forests with some soil variables changed in a favourable direction and some in an unfavourable direction.