Long-term vegetational history of a Picea abies stand in south-eastern Norway: Implications for the conservation of biological values

The development of a forest stand in south-eastern Norway during the last 9000 years is investigated by pollen and charcoal analyses. The aims are to identify factors that have influenced current biodiversity, which includes the lichen Usnea longissima, and examine the immigration and establishment of the current dominant tree Picea abies. Fire has been a variable but major disturbance factor at the site throughout the study period but has ceased during the last 100 years. Picea immigration was primarily a natural process but local establishment occurred after a major disturbance. Cultural impact began 3500 years ago during the Bronze Age, but was most intensive between AD 1600 and AD 1900. It led to the local extinction of deciduous trees and created a biodiversity bottleneck that facilitated the rise to dominance of Picea. Guidelines are proposed for future management designed to preserve and enhance local biological values.     

Artificial dispersal of endangered epiphytic lichens: a tool for conservation in boreal forest landscapes

Due to increasing fragmentation of the boreal forests of Fennoscandia, a number of epiphytic lichens are now becoming threatened. Since these species typically are limited by a poor ability of dispersal, one possible but largely unexplored strategy for conservation is to disperse lichen material artificially into suitable habitats. Therefore, the objective of this study was to evaluate survival and vitality in lichen fragments from Evernia divaricata (L.) Ach. and Ramalina dilacerata (Hoffm.) Hoffm. after transplantation into three different stand types situated in northern Sweden, using different fragment sizes and modes of transplantation. After one year, survival ranged between 85% and 97.5%, and new growth occurred at all transplantation sites. The study has shown that transplantations of small fragments might constitute a resource-efficient option for establishment of new populations of endangered lichen species, or for enlarging their populations at the present sites of occurrence. In addition, the mode of transplantation was of importance for fragment vitality.     

Effects of forest fragmentation on the granivory of differently sized seeds

The Maulino forest is a unique temperate ecosystem restricted to a small range of the coast of central Chile. This forest harbors many endemic species, and is threatened due to intensive deforestation and fragmentation. Currently the Maulino forest is composed of a suite of small fragments scattered in a landscape dominated by exotic plantations. The fragmentation of the Maulino forest has resulted in a higher abundance of granivores in small forest fragments compared with continuous forest. In order to determine if fragmentation-induced changes in granivore abundance affects the granivory of different size seeds, we experimentally assessed seed predation of a large-seeded species [Nothofagus glauca (Phil.) Krasser] and a small-seeded species [Nothofagus obliqua (Mirbel) Oersted] in the edges and interior of one continuous (large) forest and three small fragments (∼3 ha) surrounded by plantations of the exotic tree Pinus radiata. To determine what kind of granivores are preying upon seeds, seeds of both species were excluded from and exposed to large and small granivores. Granivory was higher in small fragments than in continuous forest, higher in the edges than in the forest interior, and higher upon large than on small seeds. Rodents, which were more abundant in forest fragments, were the main consumers. Thus, fragmentation indeed affects granivory increasing the consumption of seeds by predators inhabiting the Maulino forest remnants or coming from the matrix. This change may affect the future structure of the tree community in forest fragments.     

Dynamics of early wolf and cougar eradication efforts in Montana: implications for conservation

Bounty records from Montana (1902-1930) were used to better understand spatial and temporal response of wolves (Canis lupus) and cougars (Puma concolor) to an eradication program. Number of payments for wolves declined from a high of 4116 in 1903 to 0 by 1928. Numbers of cougar pelts fluctuated, but generally declined from 177 in 1908 to 2 in 1930. An inverse spatial relationship existed between density of payments for wolves and cougars. Payments for wolves were greatest in the Prairie ecoregion, while payments for cougars were greatest in the Montane. The ratio of pup to adult wolves also was greatest in the Prairie ecoregion and least in the montane. If distribution and frequency of bounty payments were a reflection of the distribution and abundance of wolves, populations were highest and most productive in areas where they currently do not occur.     

Influence of single trees on spatial and temporal patterns of belowground properties in native pine forest

Linkages between forest dynamics and ecosystem processes are poorly understood and this limits our ability to adequately estimate future changes in forest ecosystems due to human-induced global change. In particular at the single tree level, our understanding of temporal and spatial changes of belowground properties during forest succession is limited. Thus, our aim was to test whether we find a spatial and temporal gradient in nutrient availability and an associated shift in microbial community structure with increasing distance and age of single trees. We found that inorganic nitrogen was less available below the crown of single trees, while soluble organic carbon (DOC) was much more abundant, in particular in the inner zone of influence, i.e. close to the stem. The fungal:bacterial PLFA ratio was greater while microbial biomass carbon (MicC) was lower below the tree crown, indicating a strong influence of trees on spatial patterns of microbial biomass and community structure. Moreover, the positive correlation between MicC and total extractable N, and the negative correlation between fungal:bacterial biomass and δ¹⁵N, suggested that the microbial biomass was N limited below the tree crown and as a consequence nutrient cycling was presumably decelerated compared to open conditions. We also found a temporal pattern of increasing surface soil C and N content with increasing tree age (up to 250 years), underlining the significant role of single trees in creating spatial and temporal heterogeneity in forests.     

Dead wood threshold values for the three-toed woodpecker presence in boreal and sub-Alpine forest

Predicting species' responses to habitat loss is a significant challenge facing conservation biologists. We examined the response of both European three-toed woodpecker subspecies Picoides tridactylus tridactylus and P. tr. alpinus to different amounts of dead wood in a boreal and a sub-Alpine coniferous forest landscape in central Sweden and Switzerland, respectively. Habitat variables were measured by fieldwork in forests with breeding woodpeckers (n=10+12) and in control forests without breeding woodpeckers (n=10+12) in the same landscape. Logistic regression analyses revealed steep thresholds for the amount of dead standing trees and the probability of three-toed woodpecker presence in both Sweden and Switzerland. The probability of the presence of three-toed woodpeckers increased from 0.10 to 0.95 when snag basal area increased from 0.6 to 1.3 m² ha⁻¹ in Switzerland and from 0.3 to 0.5 m² ha⁻¹ in central Sweden. In Switzerland, a high road network density was negatively correlated to the presence of woodpeckers (r=−0.65, p=0.0007). The higher volumes of dead wood in Switzerland, where population trends are more positive, than in central Sweden, where the population is declining, would suggest that the volumes of dead wood in managed forests in Sweden are too low to sustain three-toed woodpeckers in the long-term. In terms of management implications, we suggest a quantitative target of at least 5% of standing trees in older forests being dead over at least 100 ha large forest areas. This corresponds about to ?1.3 m² ha⁻¹ (basal area) or ?15 m³ ha⁻¹ (volume), still depending on site productivity.     

Long-term effects of experimental forest harvesting on abundance and reproductive demography of terrestrial salamanders

Both observational and experimental studies have documented drastic reductions in salamanders after forest harvesting. Yet, the amount of time until salamander populations rebound and the factors limiting recovery after harvesting are unknown. We compared the effects of six oak regeneration practices to a control, representing a disturbance gradient from no treatment to silvicultural clearcut, on the relative abundance and reproductive demography of terrestrial salamanders through 13-years post-harvest. Following the experimental disturbance, relative abundance of terrestrial salamanders in treatments that opened the canopy were significantly and persistently lower than in either untreated control stands or midstory herbicide treatments. In general, this trend persisted through 9-13 years after treatment. Changes to demography of commonly captured salamanders varied by species, but Plethodon cinereus had a greater proportion of juveniles in unharvested treatments 7-13-years post-harvest, and Desmognathus ochrophaeus had a greater proportion of juveniles and a greater number of eggs/female in unharvested treatments 1-6-years post-harvest. Population modeling of P. cinereus indicated that adult survival had the greatest elasticity of the vital rates. Further, >60 years may be needed before P. cinereus reduced by the observed decline could reach pre-harvest levels of abundance. Of the treatments with canopy disturbance, the group selection harvest had the greatest abundances of salamanders 7-13-years post-harvest, but when coupled with future stand entries, the volume of wood fiber extracted, costs of harvesting, reduced sprouting of oaks, and soil disturbances, this method may not have the best balance of ecological and economic sustainability in central Appalachian hardwood forest.     

Evaluation of carbon exchange in a boreal coniferous stand over a 10-year period: An integrated analysis based on ecosystem model simulations and eddy covariance measurements

In order to assess the capacity of the boreal forest ecosystem to intercept atmospheric carbon over a period of years, a climate-driven growth model (FinnFor, process-based) was applied to calculate the seasonal and inter-annual variability of net ecosystem CO₂ exchange (NEE) and component carbon fluxes (gross primary production - GPP and total ecosystem respiration - TER) against a 10-year (1999-2008) period of eddy covariance (EC) measurements in a Scots pine (Pinus sylvestris L.) stand in Eastern Finland. Furthermore, the role of climatic factors, leaf area index (LAI) and physiological responses of trees regarding the ecosystem carbon fixation processes were evaluated. An hourly time-step was used to simulate the carbon exchange based on measured tree/stand characteristics and meteorological input for the experimental site, and the dynamic LAI was used throughout the 10-year simulations. The model predicted well the annual course of NEE compared to the measured values for most of the years, with the development of LAI (2.4-3.3 m² m⁻², as simulated). The simulated NEE over the study period shows that, on average, 62% of the variation refers to daily and 88% to monthly measured NEE. Both modeled and measured daily NEE showed similar responses to the temperature, photosynthetically active radiation and vapor pressure deficit during the growing seasons. In the simulation, the annual amount of GPP varied from 720.8 to 910.4 g C m⁻² with a mean value of 825.3 g C m⁻², and the annual mean TER/GPP ratio was 0.79, close to the measured value. Carbon efflux from the forest floor was the dominant contributor to the forest ecosystem respiration. The inter-annual variation of GPP mostly corresponded to the development of LAI, temperature sum and total incoming radiation over the 10-year simulation period. It was suggested that the process-based model could be applied to study the carbon processes for natural and management-induced dynamics of Scots pine forest ecosystem over longer periods across a wider climate gradient in the boreal zone.     

Population structure and movements of a threatened butterfly (Lopinga achine) in a fragmented landscape in Sweden

The red-listed butterfly Lopinga achine was studied by mark-recapture methods in southern Sweden for three seasons. We examined movement within and between populations and egg production in relation to age. The majority of the movements were small with mean movements between recaptures of 45-54 m for males and 94-116 m for females. There were few movements between sites, 20 of 996 recaptured males moved and 36 of 391 recaptured females, even though the distance to other sites was in many cases <100 m. The distance moved and the number of females moving between sites increased with increasing age. On average, a female that moves does so after laying two-thirds of its eggs in its natal site. It is therefore important to take account of the proportion of reproductive effort involved in dispersal when estimating colonisation ability. The males did not move more with increasing age. Female behaviour can be seen as a “spread-the-risk” strategy, an adaptation to the successional habitat of L. achine, whose natal site sooner or later will deteriorate. Butterflies like L. achine living in successional habitats may exhibit mobility that is intermediate between butterflies living in ephemeral habitats (very mobile) and in long-lived habitats (sedentary).     

Strong negative effect of alien herbivores on endemic legumes of the Canary pine forest

We studied the effects of herbivores on populations of four endemic legume species in pine forests in Caldera de Taburiente National Park on La Palma, Canary Islands. Seeds of these species were sown in control and herbivore-exclusion plots. Over the subsequent 4-years period, we assessed the effects of herbivores by counting the number of individual plants that emerged and by following their growth. We assessed growth over time using three plant-size categories. For these four species, we conclude that the presence of herbivores, including barbary sheep (Ammotraguslervia), goat (Caprahircus) and European rabbit (Oryctolaguscuniculus), exerts a strong negative effect on plant establishment. We suggest that the paucity of understory plant species in the pine forests of the Canary Islands may be strongly correlated with the presence of introduced herbivores in this region. We propose urgent conservation measures, such as large fenced areas, control activities and the most effective measure, eradication.     

Effects of acidification and liming on feeding groups of nematodes in coniferous forest soils

Summary Nematodes were sampled in untreated, acidified, and limed plots in a Norway spruce (Fexboda) and a Scots pine (Norrliden) stand. At Fexboda, the total number of nematodes was significantly reduced after the acidification. This reduction was probably due to a shock effect, because the samples were taken only 5 months after an application of 200 kg H₂SO₄ ha^-1 to the forest floor. However, the root/fungal-feeding Aphelenchoides was not reduced, probably because it is more tolerant of high acid concentrations than most other nematodes. At Norrliden, where the samples were taken 7 years after the last application of H₂SO₄, no significant differences were found between the acidified and untreated plots. If the treatment with H₂SO₄ caused similar effects as at Fexboda, the results indicate a recovery of the nematode populations. Decreased predation from lumbricids rather than a recovery of microfloral populations probably allowed this recovery. No marked effect of lime, spread 2 (Fexboda) and 12 years (Norrliden) before the sampling on the numbers of any of the nematode feeding groups was found. This correlated with almost no change in bacterial biomass after liming, while the active fraction of fungal hyphae was unaffected by liming at Fexboda and reduced by liming at Norrliden. A tendency for decreasing numbers of all nematode feeding groups in the limed plots at Norrliden coincided with increasing numbers of lumbricids.     

Long-term impacts of even-aged timber management on abundance and body condition of terrestrial amphibians in Northwestern California

Conservation needs for amphibians in managed timberlands may differ based upon the species present and the timber harvesting methods employed. Clearcuts have been documented to be detrimental to amphibians but the impacts of associated silvicultural edges and alternative harvesting treatments are not well understood. The primary objective of this study was to determine if amphibian abundances and body condition differed in thinned forests and intact forests, and in clearcuts and associated silvicultural edges. We also examined which environmental attributes were important in explaining observed differences. We sampled clearcuts, silvicultural edges, and adjacent late-seral forests at 10 sites in northwestern California from October 1999 to July 2002. Clearcuts at these sites ranged in age from 6 to 25 years. Five of these forest stands were intact and five had been commercially thinned at least 10 years prior to our study. Amphibian abundances were similar in thinned and unthinned forests, but body condition of the most common species was lower in thinned forests. Abundances of amphibians were nearly twice as high in forests and at silvicultural edges than in clearcuts. Clearcutting at these sites appears to have affected amphibian numbers up to 25 years post-harvest, however, silvicultural edges were suitable habitats for amphibians. While commercial thinning did not reduce amphibian numbers, it is an intermediate treatment followed by clearcutting. Where conservation of amphibians is a concern, even-aged silvicultural systems may not provide the most appropriate method for maintaining viable populations on managed forestlands in the northwestern US.     

Removal of logging residue in Norway spruce thinning stands: Long-term changes in organic layer properties

The aim of this study was to determine whether repeated removal of logging residue in Norway spruce thinning stands causes consistent long-term effects on soil microbial processes in C and N cycling and on soil concentrations of two major groups of plant secondary compounds, phenolic compounds and terpenes. The study sites were four 47-to 70-year-old Norway spruce (Picea abies (L.) Karst) stands, all growing on relatively fertile sites in southern and central Finland. These stands had been thinned twice, the first thinning 22-29 years ago and the second thinning 10 years after the first, 12-19 years ago. After thinnings, the logging residue was either removed (whole-tree harvest, WTH) or left on the site (stem-only harvest, SOH). In one experiment there was also a treatment where a double amount of logging residue was distributed on the plot. Samples were taken from the organic layer (O_fh). Removal of logging residue did not affect the C-to-N ratio or pH much. Removal of logging residue did not affect the amounts of C and N in the microbial biomass but tended to decrease the rates of net N and C mineralization (CO₂ production). It decreased both sesquiterpene and diterpene (mostly resin acids) concentrations but did not affect triterpene (mostly sterols) concentrations. Neither total water-soluble phenolic compounds nor an important group of phenolic compounds, condensed tannins, were affected by removal of logging residue. At these four study sites, the effects of removing logging residue were strongest on the two least fertile sites, whereas the most fertile site did not respond much to removal. In conclusion, with regard to the processes and ratios indicating N availability, stem-only harvest seems generally to be more favorable than whole-tree harvest in long-term.     

Microsatellite analysis of genetic population structure of the endangered cyprinid Anaecypris hispanica in Portugal: implications for conservation

The endangered fish species Anaecypris hispanica is restricted to eight disjunct populations in the Portuguese Guadiana drainage. The genetic structure of these populations was studied in order to determine levels of genetic variation within and among populations and suggest implications for conservation of the species. Based on five microsatellite loci, the null hypothesis of population homogeneity was tested. Tests for genetic differentiation revealed highly significant differences for pairwise comparisons between all populations, and substantial overall population subdivision (F_ST=0.112). All sampled populations contained unique alleles. Our findings indicate marked genetic structuring and emphasise limited dispersal ability. The high levels of genetic diversity detected within and among A. hispanica populations suggest, however, that the observed fragmentation and reduction in population size of some populations during the last two decades, has impacted little on levels of genetic variability. Data imply that most A. hispanica populations should be managed as distinct units and that each has a high conservation value containing unique genetic variation. It is argued that geographic patterns of genetic structuring indicate the existence of eight management units.     

Island differences in population size structure and catch per unit effort and their conservation implications for Komodo dragons

Species inhabiting archipelagos are often characterised by high levels of interpopulation divergence (e.g. size related traits). This divergence may, in turn, influence their life-history. To facilitate better management and conservation of the Komodo dragon (Varanus komodoensis), an island endemic, we identified demographic differences between two island populations in Komodo National Park, Indonesia. Comparison of data collected from dragon populations inhabiting Rinca Island and the much smaller Gili Motang Island indicated that between 1994 and 2004, the Komodo dragon population on Gili Motang significantly decreased its: (1) mean body mass, (2) body condition and (3) relative abundance. These results suggest that the numerically small Gili Motang population was oscillating downwards; in contrast, the Rinca Island population had been relatively stable. More importantly these results emphasize the necessity for managers of this priority conservation species to understand further the inherent functional differences among dragon populations to develop island specific management units. Current management practices (e.g. monitoring) instigated by Komodo National Park management ignore small island dragon populations and thus run the risk of being unable to detect adverse effects for populations that are potentially most prone to decline.     

Effects of soil application of olive mill wastewaters on the structure and function of the community of arbuscular mycorrhizal fungi

Recycling of olive mill wastewaters (OMW) into agricultural soils is a controversial issue since benefits to soil fertility should counterbalance potential short-term toxicity effects. We investigated the short-term effects of OMW on the soil-plant system, regarding the diversity, structure and root colonization capacity of arbuscular mycorrhizal (AM) fungi and the respective growth response of Vicia faba L, commonly used as green manure in olive-tree plantations. A compartmentalized pot system was used that allowed the establishment of an AM fungal community in one compartment (feeder) and the application of three OMW dose levels in an adjacent second compartment (receiver). At 0, 10, and 30 days after OMW treatment (DAT), V. faba pre-germinated seeds were seeded in the receiver compartment. At harvest, shoot and root dry weights, AM fungal root colonization, soil hyphal length and P availability were recorded in the receiver compartment. In addition, OMW effects on AM fungal diversity in plant roots were studied by DGGE. A transient effect of OMW application was observed; plant growth and AM fungal colonization were initially inhibited, whereas soil hyphal length was stimulated, but in most cases differences were absent when seeding was performed 30 DAT. Similarly, changes induced in the structure of the root AM fungal community were of transient nature. Cloning and sequencing of all the major DGGE bands showed that roots were colonized by Glomus spp. The transient effects of OMW on the structure and function of AM fungi could be attributed to OMW-derived phytoxicity to V. faba plants or to an indirect effect via alteration of soil nutritional status. The high OMW dose significantly increased soil P availability in the presence of AM fungi, suggesting efficient involvement of AM fungi in organic-P minerilization. Overall our results indicate that soil application of OMW would cause transient changes in the AM fungal colonization of V. faba plants, which, would not impair their long-term plant growth promoting ability.     

An unseen foe in arthropod conservation efforts: The case of Wolbachia infections in the Karner blue butterfly

Infection by endosymbiotic bacteria is an underappreciated threat to endangered arthropods with serious implications for captive management programs. We examined the nature of Wolbachia infection in the North American endangered Karner blue butterfly, Lycaeides melissa samuelis, as a case study. Screening for Wolbachia across the range of the species confirmed widespread infection in the western half of the Karner blue’s range. Multilocus sequence typing using six genes confirmed that the infection in the western populations is attributable to a single strain of Wolbachia. This strain was also detected in the closely related Melissa blue butterfly, L. m. melissa, the presumed source of the infection. The infection in the Karner blue butterfly was perfectly correlated with the presence of a foreign mitochondrial DNA variant present in the Melissa blue butterfly, consistent with the hypothesis that the mitochondrial introgression was driven by the spread of Wolbachia. A single individual out of 71 screened from the eastern portion of the range of the Karner blue butterfly was also infected, however this infection was attributable to a different strain. Simulation models of the demographic effects of the spread of Wolbachia infection to uninfected populations and metapopulations suggest that such an infection might further reduce already small population sizes and substantially increase the probability of population extirpation. We discuss threats to other endangered arthropods in the light of this case study and make recommendations for minimizing the impact of endosymbiont infections in conservation plans, especially those including captive propagation and augmentation of endangered arthropod populations.     

Land-use history and plant performance in populations of Trillium grandiflorum

Many forest herbs show reduced frequency in forests growing on former agricultural land (secondary forests) relative to forests that were never cleared for agriculture (primary forests). To explain this pattern, studies of secondary forest colonization by forest herbs have stressed the role of limited seed dispersal, though environmental conditions may also limit colonization. Here I demonstrate that adult plant performance of Trillium grandiflorum is significantly reduced in secondary vs. primary forests in central New York State, USA. Comparing primary and secondary sites both with relatively high soil pH, T. grandiflorum populations were more dense, and individual plants were larger and more likely to be flowering in primary than secondary forests. Using counts of annual leaf scars on the rhizome, I demonstrate that this result is not due to secondary populations being younger than primary populations. Age-specific leaf area was significantly greater in primary than secondary stands at high soil pH, and the data suggest slightly reduced seed set in secondary stands as well. These results point to a more important role for environmental conditions in limiting forest-herb colonization of secondary forests than has been previously appreciated. Restoration and conservation of forest herbs in post-agricultural forests will require more than simply overcoming dispersal limitation.     

Strong bottom-up and weak top-down effects in soil: nematode-parasitized insects and nematode-trapping fungi

The soils of the Bodega Marine Reserve (BMR, Sonoma County, California) contain many nematode-trapping fungi and many ghost moth larvae parasitized by entomopathogenic nematodes. The current study determined whether these nematode-parasitized moth larvae, which can produce very large numbers of nematodes, enhanced the population densities of nematode-trapping fungi and whether the fungi trapped substantial numbers of nematodes emerging and dispersing from moths. Wax moths were used in place of ghost moths because the former are easier to obtain. When nematode-parasitized moth larvae were added to laboratory microcosms containing BMR field soil, the population densities of four nematode-trapping fungi increased substantially. The greatest increase in population density was by Arthrobotrys oligospora, which uses adhesive networks to capture nematodes. A. oligospora population density increased about 10 times when the added moth larvae were parasitized by the nematode Heterorhabditis marelatus and about 100 times when added moth larvae were parasitized by the nematode Steinernema glaseri. Other trapping fungi endemic to the soil and enhanced by nematode-parasitized moth larvae included Myzocytium glutinosporum, Drechslerella brochopaga, and Gamsylella gephyropaga, which produce adhesive spores, constricting rings, and adhesive branches, respectively. The data suggest that the previously documented abundance and diversity of nematode-trapping fungi in BMR soil can be explained, at least in part, by nematode-parasitized insects, although that inference requires further studies with ghost moths. The strong bottom-up enhancement of nematode-trapping fungi was not matched by a strong top-down suppression of nematodes, i.e. the fungi trapped fewer than 30% of dispersing nematodes.     

Linking forest structure and composition: avian diversity in successional forests of Chiloé Island, Chile

We show how Chilean forest bird species richness, abundance and guild structure changes as a function of structural properties of forest stands. We surveyed bird assemblages in two old-growth (>200 years), two mid-successional (30-60 years), and two early-successional forest stands (4-20 years), from November 1999 to September 2000, on Chiloé Island, southern Chile (42°S). Birds were grouped into four habitat-use guilds: large-tree users, vertical-profile generalists, understory species, and shrub-users that occasionally use forests. We recorded a total of 24 bird species: 21 in old-growth, 14 in mid-successional and 16 in early-successional stands. Large-tree users and understory birds were most abundant in old-growth stands, vertical-profile generalists were common in both old-growth and mid-successional stands, and shrub-users were only common in early-successional stands. For nine bird species we found significant relationships between their local abundance and forest structural elements. Higher bird densities in old-growth forests were associated with greater availability of canopy emergent trees, snags, logs and understory bamboo cover in this habitat. Accordingly, bird species diversity in forest stands can be predicted by the presence of these structural elements, and forests should be managed to conserve structural elements that create favorable habitat for bird species in order to prevent future species losses due to logging practices.