Influence of variable retention harvests on forest ecosystems: Plant and mammal responses up to 8 years post-harvest

Green-tree retention systems are an important management component of variable retention harvests in temperate zone coniferous forests. Residual live trees (“legacy trees”) provide mature forest habitat, increase structural diversity, and provide continuity in the regenerating stand. This study was designed to test the hypotheses that, at up to 8 years after harvest, abundance and species diversity of communities of (i) understory plants and (ii) forest-floor small mammals, and (iii) relative habitat use by mule deer (Odocoileus hemionus), will decline with decreasing levels of tree retention. Communities of plants and forest floor small mammals were sampled in replicated clearcut, single seed-tree, group seed-tree, patch cut, and uncut forest sites in mixed Douglas-fir (Pseudotsuga menziesii)—lodgepole pine (Pinus contorta) forest in southern British Columbia, Canada from 2000 to 2003 (5–8 years post-harvest). Habitat use by mule deer was measured during summer and winter periods each year from 1999 to 2003 in these same sites.     

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.     

Distribution and abundance of small rodents in Croatian forests

The relative abundance and the population development of small rodents were investigated on 17 sites in Croatian forests from September 1999 through June 2003. Common snap traps were set for two or three nights along transect lines in different forest types. In 27,598 trap–nights, 2,151 specimens were trapped. Small rodents were present in up to 45% of the traps in pedunculate oak forests dominated by Quercus robur L. In common beech forests with a predominance of Fagus sylvatica L., the percentage of occupied traps reached 59%. At most sites the following species were dominant:Clethrionomys glareolus Schr., Apodemus agrarius Pall. and A. flavicollis Melch. Additionally, in a forest community of Carpino betuli-Quercetum roboris typicum R., the absolute number of small rodents was calculated by means of the standard minimum method. The minimum rodent density was observed in April 2003 with 11.63 n/ha (number of specimens per hectare). The maximum was reached in March 2001 with 82.03 n/ha. Near the same site, the absolute number of the small rodent population was estimated using the Y method in spring 2001 as approximately 101.34 n/ha. In general, the abundance of small rodents was dependent on the forest seed crop.     

Forecasting vole population outbreaks in forest plantations: The rise and fall of a major mammalian pest

Voles of the genera Microtus and Myodes feed on tree seedlings planted on cutover forest land in temperate and boreal forests of North America and Eurasia. This damage may have serious economic implications as well as limit regeneration of appropriate tree species in certain forest ecosystems. Prediction of vole population outbreaks and feeding damage to forest plantations, across even a limited geographic range, has yet to be achieved in North America. Thus, a major objective was a detailed analysis of changes in population dynamics of long-tailed voles (Microtus longicaudus), and to test three hypotheses (H) that vole populations would: (H₁) rise and fall in accordance with the abundance of herbaceous plants (grasses and forbs) during early vegetative succession after forest harvesting, (H₂) be positively associated with grass-seeded sites; and (H₃) incidence of feeding damage to seedlings would be positively associated with vole abundance. Voles were live-trapped for 6 years (2004–2009) from the time of harvesting on intensive sites, as well as surveyed over a range of extensive sites. Population numbers were related to habitat characteristics and tree damage in young forest plantations near Golden, British Columbia, Canada.     

Site, plot, and individual tree yield reduction of interior Douglas-fir associated with non-lethal infection by Armillaria root disease in southern British Columbia

Root pathogens are one of the principle factors affecting forest productivity in many forests, but few estimates of impact are available. Non-lethal root infections associated with Armillaria root disease were studied to determine their effect on stem volume yield in seven planted Douglas-fir stands and a naturally regenerated stand in British Columbia's southern interior. Trees were removed from the soil and the infection date of a random selection of trees was determined. The volume reduction attributable to disease was determined as a comparison of diseased to disease-free trees over time since infection. Volume reductions per tree ranged from 0 to 30 dm³ (0-27%) depending on the tree age and disease duration. Yield reduction reached 27 m³/ha, averaging 15 m³/ha for the three oldest planted sites by age 30 (7-15%), but was lower at the naturally regenerated site. Yield reduction at the site level correlated best with the number of diseased trees and an unknown site factor. Sites with slow juvenile growth had the least yield reduction owing to their lower incidence of disease over time. Yield was less affected by the proportion of diseased primary roots per tree than by the cumulative time since infection. A few of the diseased trees maintained growth rate after infection similar to disease-free trees; interestingly, these trees were smaller than average to begin with. Overall, trees suffer accumulating growth reduction without recovery. Root diseases prevent full expression of site potential even without mortality. Minimizing disease impact in respect to other forest management goals is also discussed.     

Red-tree vole habitat suitability modeling: Implications for conservation and management

In the Pacific Northwest, USA, red-tree voles (Arborimus longicaudus) are of conservation and management interest owing to their apparent association with late-seral forests and the relatively small extent of such forests, largely a function of timber harvest, fire, and conversion of forests to non-forest uses during the past century. We created and evaluated a series of red-tree vole habitat association models, and applied the best model to evaluate tree vole habitat quality within and outside of reserves throughout most of their range in Oregon and northern California. We modeled presence and absence of tree vole nests across a gradient of biotic, abiotic, and spatial features; and within and outside of reserves. The best model included spatial coordinates, percent slope, basal area of trees with diameter at breast height (dbh) between 45 and 90 cm, maximum tree dbh, and standard deviation of conifer dbh. Plots with tree vole nests contained many late-seral/old-growth forest attributes such as large diameter, older, and variably sized trees. Evaluation of the best model, including rigorous cross-validation, showed the model to be statistically robust and to have very good/excellent predictive ability. Reserves had significantly higher mean habitat quality than non-reserved lands, and reserves had much more high quality habitat than non-reserves.     

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.     

Forest stand dynamics and sudden oak death: Mortality in mixed-evergreen forests dominated by coast live oak

Sudden oak death (SOD), caused by the recently discovered non-native invasive pathogen, Phytophthora ramorum, has already killed tens of thousands of native coast live oak and tanoak trees in California. Little is known of potential short and long term impacts of this novel plant–pathogen interaction on forest structure and composition. Coast live oak (Quercus agrifolia) and bay laurel (Umbellularia californica) form mixed-evergreen forests along the northern California coast. This study measured tree mortality over a gradient of disease in three time periods. Direct measurements of current mortality were taken during 2004, representing a point-in-time estimate of present and ongoing mortality. Past stand conditions, c. 1994, were estimated using a stand reconstruction technique. Future stand conditions, c. 2014, were calculated by assuming that, given a lack of host resistance, live trees showing signs of the disease in 2004 would die. Results indicate that coast live oaks died at a rate of 4.4–5.5% year⁻¹ between 1994 and 2004 in highly impacted sites, compared with a background rate of 0.49% year⁻¹, a ten-fold increase in mortality. From 2004 to 2014, mortality rates in the same sites were 0.8–2.6% year⁻¹. Over the entire period, in highly impacted sites, a 59–70% loss of coast live oak basal area was predicted, and coast live oak decreased from 60% to 40% of total stand basal area, while bay laurel increased from 22% to 37%. Future stand structures will likely have greater proportions of bay laurel relative to coast live oak.     

Presettlement and modern disturbance regimes in coast redwood forests: Implications for the conservation of old-growth stands

Coast redwood (Sequoia sempervirens), a western North American conifer of ancient lineage, has a paradoxical combination of late-successional characteristics and strong adaptations to disturbance. Despite its shade tolerance and heavy dominance of the canopy on many sites, redwood saplings are uncommon in upland old-growth stands. Information needed to ensure the conservation of old-growth redwood forests has been limited. In this review paper, we integrate evidence on redwood biology with data on the historic and modern disturbance regimes to help clarify the degree to which key attributes of redwood forests may have been dependent upon periodic disturbance. Available evidence suggests that episodes of fire, flooding, and slope failure prior to European settlement were frequent but predominantly of low to moderate severity and extent, resulting in broadly uneven-aged forests. The majority of fires prior to European settlement were apparently of human origin. Frequency and severity of the major disturbance agents have been radically changed in modern times. Fires have been largely excluded, and flooding has been altered in ways that have often been detrimental to old-growth redwoods on alluvial terraces. However, because of the apparent anthropogenic origin of most presettlement fires, the long-term evolutionary role of fire for coast redwood is ecologically ambiguous. With fire exclusion, redwood possibly could be displaced to some extent on upland sites by increasing abundance of fire-sensitive competitors. Alternatively, redwood may be able to maintain dominance by vegetative sprouting and new seedling establishment on root-wad mounds, fallen logs, and on soil exposed by slope failure. Future research priorities are suggested that will help resolve some of the current ambiguities.     

Local variability of stand structural features in beech dominated natural forests of Central Europe: Implications for sampling

Current knowledge of the within-site variability of major stand structural features in beech dominated natural forests is limited. Numerous studies have used just several small plots for characterizations of natural stands, but this may lead to generalizations based on unreliable results. This study shows how major stand structural features vary at the local scale, and how suitable sampling may reflect this variability. Stem position maps of three natural forests in the Czech Republic (Zofin 71 ha, Salajka 19 ha and Zakova hora 17 ha) were used. Each vector stem position map representing all live and dead trees with DBH ≥ 10 cm was intensively analyzed using square sample plots of different sizes (10 × 10 m; 20 × 20 m; 30 × 30 m; 50 × 50 m; 100 × 100 m; 140 × 140 m and 200 × 200 m). Basic statistics (mean, standard deviation, coefficient of variation, min., and max.) were calculated for every plot size and each of six major stand features: density, basal area and volume of living trees, volume of course woody debris, total volume and proportion of course woody debris in total volume.     

Predisposition to bark beetle attack by root herbivores and associated pathogens: Roles in forest decline,gap formation,and persistence of endemic bark beetle populations

Bark beetles are largely known for their ability to undergo intermittent population eruptions that transform entire landscapes and pose significant economic hardships. However, most species do not undergo outbreaks, and eruptive species usually exert only minor disturbances. Understanding the dynamics of tree-killing noneruptive species can provide insights into how beetles persist at low densities, and how some spatiotemporal patterns of host predisposition may more likely favor breaching eruptive thresholds than others. Elucidating mechanisms behind low-density populations is challenging, however, due to the requirement of long-term monitoring and high degrees of spatial and temporal covariance. We censused more than 2700 trees annually over 7 years, and at the end of 17 years, in a mature red pine plantation. Trees were measured for the presence of bark beetles and wood borers that breed within the primary stem, root weevils that breed in root collars, and bark beetles that breed in basal stems. We quantify the sequence of events that drive this decline syndrome, with the primary emergent pattern being an interaction between below- and above-ground herbivores and their fungal symbionts. This interaction results in an expanding forest gap, with subsequent colonization by early-successional vegetation. Spatial position strongly affects the likelihood of tree mortality. A red pine is initially very likely to avoid attack by tree-killing Ips beetles, but attack becomes increasingly likely as the belowground complex spreads to neighboring trees and eventually make trees susceptible. This system is largely internally driven, as there are strong gap edge, but not stand-edge, effects. Additional stressors, such as drought, can provide an intermittent source of susceptible trees to Ips beetles, and elevated temperature slightly accentuates this effect. New gaps can arise from such trees as they subsequently become epicenters for the full complex of organisms associated with this decline, but this is not common. As Ips populations rise, there is some element of positive feedback, in that the proportion of killed trees that were not first colonized by root organisms increases. This positive feedback is very weak, however, and we propose the slope between beetle population density and reliance on host stress as a quantitative distinction along a gradient from noneruptive through eruptive species. Almost all trees colonized by Ips were subsequently colonized by wood borers, likely a source of negative feedback. We discuss implications to our overall understanding of cross-scale interactions, between-guild interactions, forest declines, and eruptive thresholds.     

Diversity and structure of regenerating tropical dry forests in Costa Rica: Geographic patterns and environmental drivers

Much of the dry tropical forest biome has been converted to agricultural land uses over the past several centuries. However, in conservation areas such as those in the Guanacaste and Tempisque regions of Costa Rica, tropical dry forests are regenerating due to management practices including fire suppression. To better understand the patterns of secondary succession occurring in Costa Rican tropical dry forest, we established 60 20 × 50 m plots in mature and regenerating forests in the Sector Santa Rosa (formerly known as Parque Nacional Santa Rosa) and Palo Verde National Park. Plots were stratified into three plant communities: tropical dry oak forest (Quercus oleoides) (SROAK), Santa Rosa tropical dry forest (SRTDF), and Palo Verde tropical dry forest (PVTDF). In these plots we measured and identified and all individuals >10 cm DBH, measured but did not identify stems <10 cm but taller than 1.3 m, counted woody seedlings (<1.3 m height) and analyzed soil chemical and physical properties.     

Historical fire and vegetation dynamics in dry forests of the interior Pacific Northwest,USA, and relationships to Northern Spotted Owl (Strix occidentalis caurina) habitat conservation

Regional conservation planning frequently relies on general assumptions about historical disturbance regimes to inform decisions about landscape restoration, reserve allocations, and landscape management. Spatially explicit simulations of landscape dynamics provide quantitative estimates of landscape structure and allow for the testing of alternative scenarios. We used a landscape fire succession model to estimate the historical range of variability of vegetation and fire in a dry forest landscape (size ca. 7900 km²) where the present-day risk of high severity fire threatens the persistence of older closed canopy forest which may serve as Northern Spotted Owl (Strix occidentalis caurina) habitat. Our results indicated that historically, older forest may have comprised the largest percentage of the landscape (∼35%), followed by early successional forest (∼25%), with about 9% of the landscape in a closed canopy older forest condition. The amount and condition of older forest varied by potential vegetation type and land use allocation type. Vegetation successional stages had fine-grained spatial heterogeneity in patch characteristics, with older forest tending to have the largest patch sizes among the successional stages. Increasing fire severities posed a greater risk to Northern Spotted Owl habitat than increasing fire sizes or frequencies under historical fire regimes. Improved understanding of historical landscape-specific fire and vegetation conditions and their variability can assist forest managers to promote landscape resilience and increases of older forest, in dry forests with restricted amounts of habitat for sensitive species.     

Effects of litter quality and parent material on organic matter characteristics and N-dynamics in Luxembourg beech and hornbeam forests

To test effects of litter quality and soil conditions on N-dynamics, we selected seven forests in Luxembourg dominated by beech (Fagus sylvatica, L.) and hornbeam (Carpinus betulus L.), and located on acid loam, decalcified marl or limestone, and measured organic matter characteristics, microbial C and N and net N-mineralization in a laboratory incubation experiment. Organic layer characteristics were significantly affected by species, with lower litter decay and higher accumulation under the less palatable beech, even on limestone. However, beech and hornbeam did not show any differences in N-cycling at all. Instead of species, N-cycling was affected by site conditions, albeit different than expected. Microbial N generally increased from acid loam to limestone, but acid loam showed higher net N-mineralization, especially in the organic layer. Also, acid loam showed high instead of low efficiency of N-mineralization per unit microbe, in both organic layer and mineral topsoil. In addition, acid loam showed net consumption of DOC instead of release in both soil layers, which suggests that not N, but C was a limiting factor to decomposition. In contrast, limestone showed low net N-mineralization in the organic layer, despite high mass and well-decomposed organic matter, and low efficiency of N-mineralization per unit microbe in both organic layer and mineral topsoil. DOC was net released instead of consumed, which supported that not C, but N was a limiting factor. The general lack of differences in net N-cycling between species, but relatively clear site effects, is discussed in relation to different microbial strategies. Acid soil may have high net N-release despite low biological activity, because N-requirements of fungi are also low, while in calcareous soil, high bacterial N-demand may counteract high gross N-release. Thus, species producing litter that decomposes rapidly may be planted to improve soil conditions and plant biodiversity, but litter quality effects on N-availability may be less important than soil conditions.     

Distribution and nest site selection of red-shouldered hawks (Buteo lineatus) in forests of northeastern Wisconsin (USA)

Red-shouldered hawks (Buteo lineatus) are threatened in Wisconsin and when nest sites are found during the cruising or marking stage of timber harvesting, the harvest is altered to accommodate the hawks. If nest site locations are known before initiation of timber harvest, foresters can employ a proactive approach to manage red-shouldered hawks while maintaining timber production. We searched for red-shouldered hawks nest sites on Marinette County Forest (MCF) which encompasses 94,000 ha in northeastern Wisconsin and is the second largest county forest in the state. We used a comparative modeling approach to evaluate distribution and habitat relationships of red-shouldered hawk nest sites in relation to a suite of environmental variables in MCF. Models were used to develop forest management recommendations for red-shouldered hawks in Wisconsin. During the spring of 2006 and 2007, we broadcasted conspecific calls to survey 1121 calling stations along forest roads and trails. We located 20 and 25 active nesting territories in 2006 and 2007, respectively (11 of which were active in both years). To understand nest site selection, we measured 22 habitat variables within 0.04-ha plots at active nest sites (n = 34) and at stratified random sites (n = 61). Logistic regression with information-theoretic model selection identified a model including greater tree species richness and closeness to forested wetland as the best-approximating model. Variable selection with Discriminant Function Analysis (DFA) indicated that nest selection was best explained by greater number of tree species, closer distance to forested wetlands, greater volume of downed woody debris, fewer small sawlogs, and increased proximity to streams. Univariate comparisons identified four of the five aforementioned variables in the DFA model as significant. Red-shouldered hawks are likely more common in Wisconsin than their state status suggests. Forest management for red-shouldered hawk nest sites should focus on increasing tree species richness, increasing down woody debris volume, and protecting forested wetlands. These recommendations may assist property managers to locate and plan for continued persistence of this species on MCF.     

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.     

Reconstructing forest history from archaeological data: A case study in the Duero basin assessing the origin of controversial forests and the loss of tree populations of great biogeographical interest

The study of more than 700 charcoals from the Vaccaean settlement of Pintia (Padilla del Duero, Valladolid) provides new local and detailed data about the Iron Age II forest composition in Continental-Mediterranean inland Spain. Pinus gr. sylvestris/nigra (from which an unexpectedly high number of charcoals were identified), together with Mediterranean pines (also highly represented), Juniperus L. and evergreen and deciduous Quercus, were the main taxa living in the area, revealing two different marked biogeographical elements in the late-Holocene landscapes of the Duero basin: Mediterranean and Eurosiberian. On one hand, the results shed new light on the assessment of the origin of Mediterranean pinewoods in the area. On the other hand, the charcoal assemblage of Pintia reveals a more important presence of the current Mediterranean highland pines in the lowlands of the Central Iberian Peninsula in the late-Holocene suggesting the existence of interglacial refugia. Finally, this work highlights palaeoecological studies as valuable tools to support decisions on forest management related to conservation and restoration of our natural heritage.     

Characteristics of aspen infected with heartrot: Implications for cavity-nesting birds

Phellinus tremulae is an important fungal decay agent common to aspen and a critical component to the cavity-nesting bird complex found in western aspen stands. Little information exists on the conditions that facilitate infection and spread of P. tremulae in aspen forests. I used Forest Inventory and Analysis (FIA) data to explore the relationships of several tree and stand characteristics to the presence and frequency of P. tremulae in aspen measured across several western states of the United States. Results suggest a strong relationship between tree age, tree diameter, and compacted crown ratio with infection frequency in trees while stand purity, canopy cover and stand age had a positive relationship with the occurrence of P. tremulae in forest stands containing aspen. Logistic regression modeling identified stand age as the only variable that increased the odds of predicting infection at the stand-level while all tree-level variables were included in the tree model. Data also show that infection rates in the study area were lower than in other parts of aspen's range, and that average size of infected trees was smaller in the study area than those reported elsewhere. These results have important implications to management of aspen for wildlife, especially for birds that use decayed aspen for nesting.     

Post-logging regeneration and recruitment of shihuahuaco (Dipteryx spp.) in Peruvian Amazonia: Implications for management

Over the past decade, shihuahuaco timber - comprising several species of Dipteryx (Fabaceae) traded internationally as “cumarú” or “Brazilian teak” - has become one of the most highly demanded types of hardwood from Peruvian Amazonia, particularly in the Chinese market. Dipteryx is an ecologically important canopy-emergent genus with widespread distribution in neotropical forests. To assess the response of Dipteryx to logging, we conducted inventories in three logging areas in the Regions of Ucayali and Loreto, Peru. The size-class distributions of Dipteryx populations in recently logged sites showed that initial post-logging conditions enhance recruitment of residual seedlings. These conditions are created by a combination of logging gaps and the activities of farmers migrating into logged lands. Through protection and liberation of shihuahuaco seedlings in post-logged forest as well as within and around agricultural fields, local residents and timber companies could favor the recovery of this valued resource. However, as logged land is increasingly converted to agriculture and pastureland, the reestablishment of mature seed trees is not assured.