Nitrogen dynamics across silvicultural canopy gaps in young forests of western Oregon

Silvicultural canopy gaps are emerging as an alternative management tool to accelerate development of complex forest structure in young, even-aged forests of the Pacific Northwest. The effect of gap creation on available nitrogen (N) is of concern to managers because N is often a limiting nutrient in Pacific Northwest forests. We investigated patterns of N availability in the forest floor and upper mineral soil (0–10 cm) across 6–8-year-old silvicultural canopy gaps in three 50–70-year-old Douglas-fir forests spanning a wide range of soil N capital in the Coast Range and Cascade Mountains of western Oregon. We used extractable ammonium (NH₄⁺) and nitrate (NO₃⁻) pools, net N mineralization and nitrification rates, and NH₄⁺ and NO₃⁻ ion exchange resin (IER) concentrations to quantify N availability along north-south transects run through the centers of 0.4 and 0.1 ha gaps. In addition, we measured several factors known to influence N availability, including litterfall, moisture, temperature, and decomposition rates. In general, gap-forest differences in N availability were more pronounced in the mineral soil than in the forest floor. Mineral soil extractable NH₄⁺ and NO₃⁻ pools, net N mineralization and nitrification rates, and NH₄⁺ and NO₃⁻ IER concentrations were all significantly elevated in gaps relative to adjacent forest, and in several cases exhibited significantly greater spatial variability in gaps than forest. Nitrogen availability along the edges of gaps more often resembled levels in the adjacent forest than in gap centers. For the majority of response variables, there were no significant differences between northern and southern transect positions, nor between 0.4 and 0.1 ha gaps. Forest floor and mineral soil gravimetric percent moisture and temperature showed few differences along transects, while litterfall carbon (C) inputs and litterfall C:N ratios in gaps were significantly lower than in the adjacent forest. Reciprocal transfer incubations of mineral soil samples between gap and forest positions revealed that soil originating from gaps had greater net nitrification rates than forest samples, regardless of incubation environment. Overall, our results suggest that increased N availability in 6–8-year-old silvicultural gaps in young western Oregon forests may be due more to the quality and quantity of litterfall inputs resulting from early-seral species colonizing gaps than by changes in temperature and moisture conditions caused by gap creation.     

Predicting canopy macrolichen diversity and abundance within old-growth inland temperate rainforests

Windward slopes of the inland mountain ranges in British Columbia support a unique inland temperate rainforest (ITR) ecosystem. Increasing fragmentation and the loss of old ITR stands have highlighted the need for determining conservation biology priorities among remaining old forest stands. We have addressed this issue by surveying foliose macrolichens within 53 old ITR stands in British Columbia's 135,000 ha very wet-cool interior-cedar hemlock (ICHvk2) biogeoclimatic subzone in the upper Fraser River watershed. Study plots were stratified by leading tree species and by “wet” versus “dry” relative soil moisture conditions. Other plot variables included: temperature, precipitation, incident solar loading, and canopy openness. Ordination plots showed a distinct assemblage of foliose cyanolichens, including Lobaria pulmonaria, Lobaria retigera, Sticta fuliginosa, Nephroma isidiosum, Nephroma occultum, and Pseudocyphellaria anomala, whose abundance was correlated with increasing relative soil moisture, temperature, canopy openness, precipitation, and basal area of spruce. Logistic regression models similarly identified relative soil moisture and temperature in all parsimonious models. Leading tree species, in combination with “wet” relative soil moisture and/or temperature, were important factors explaining the presence or absence of five (Cavernularia hultenii, L. retigera, N. occultum, Platismatia norvegica, and Sticta oroborealis) of the eight modeled old-growth associate lichen species. This combination of conditions favouring the development of canopy lichen communities in old forests was best expressed in low elevation water receiving sites. We hypothesize that groundwater availability in these sites promotes species richness and abundance of canopy lichens by creating more favourable conditions for growth, and by reducing fire return intervals which allows for the accumulation of rare species over time. Historically, forests in these wet “toe-slope” positions were disproportionately targeted for logging. Their conservation should now be a high priority, given their disproportionate significance to maintaining canopy lichen diversity in the present-day landscapes.     

Lichen diversity in temperate montane forests is influenced by forest structure more than climate

Although the effect of forest management on lichens in temperate forests has been widely examined, little is known about the influence of management-related factors on their biodiversity relative to factors that cannot be altered by management. Here we determined whether forest structure or climate determines lichen diversity in the Bavarian Forest National Park in southeastern Germany, taking spatial variables into account. We investigated 517 single tree stems along 4 transects in 113 pre-stratified plots (8 m in diameter) in this montane forest. We grouped environmental variables into three sets: climate (macroclimate, non-manageable), forest structure (manageable), and space. The explanatory powers of these sets of variables for lichen diversity were compared using variance partitioning for the lichen community, species density, and threatened species density. The relationships of single characteristics of forest structure with lichen species diversity were analyzed using generalized linear models (GLM). Lichen diversity was better explained by stand structures than by climate. Spatial effects influenced the number of species per plot. Among the structural features, the availability of dead wood and sycamore maple as well as forest continuity were most important for the enhancement of lichen diversity. Open canopy structures affected the total diversity positively. Although the availability of large trees was not an influential factor in the GLM at the plot level, high diversity levels were generally associated with large stem diameters at the level of single stems. We provide recommendations for sustainable forest-management practices that aim at specifically enhancing lichen diversity in temperate areas experiencing low levels of air pollution.     

Liana habitat and host preferences in northern temperate forests

Lianas and other climbers are important ecological and structural components of forest communities. Like other plants, their abundance in a given habitat depends on a variety of factors, such as light, soil moisture and nutrients. However, since lianas require external support, host tree characteristics also influence their distribution. Lianas are conspicuous life forms in tropical regions, but in temperate areas, where they are less prominent, little is known about factors that control their distributions in these forests. We surveyed the climbing plant species in 20 mature (100 years and greater) forested habitats in the Midwest USA at a variety of levels from simple presence/absence, to ground layer abundances, to those species that had ascended trees. We also examined attributes of the tree species with climbers attached to them. Using cluster analysis, we distinguished five different tree communities in our survey locations. We determined that 25% of the trees we surveyed had one or more lianas attached to it, with Parthenocissus quinquefolia (Virginia creeper) the most common climbing species encountered. Canopy cover and soil attributes both influenced climber species presence/absence and ground layer climber abundance. The proportion of liana species of a given climbing type (roots, stem twiner, tendril climber) was significantly related to the DBH of the host tree, with more root climbers and fewer stem and tendril climbers on large trees. In general, the DBH of climbing lianas had a significant positive relationship to the DBH of the host tree; however this varied by the identity of the liana and the tree species. The greater the DBH of the host tree, the higher the probability that it was colonized by one or more lianas, with tree species such as Pinus banksiana (jack pine) and Quercus alba (white oak) being more susceptible to liana colonization than others. Finally, some liana species such as Celastrus scandens (American bittersweet) showed a preference for certain tree species (i.e., P. banksiana) as hosts. The information obtained about the relationship between the tree and climber community in this study provides insight into some of the factors that influence liana distributions in understudied temperate forest habitats and how lianas contribute to the structure of these mature forests. In addition, these data can provide a point of comparison to other liana communities in both temperate and tropical regions.     

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.     

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.     

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.     

Dynamics of leaf area index and canopy openness of three forest types in a warm temperate zone

Deciduous broad-leaved forests (DBF), Larix principis-rupprechtii (LF) and Pinus tabulaeformis plantations (PF) are three typical forest communities in the warm temperate zone of the Dongling Mountains. In this study, we used an indirect method, hemispheric photography, to measure and analyze the dynamics of leaf area index (LAI) and canopy openness of the three forest communities. The results show that the LAI values of DBF and LF increased gradually with plant growth and development. The highest LAI value appeared in August, while canopy openness changed inversely with LAI. The lowest value appeared in November. DBF maintained a higher LAI in August and had a more open canopy in November compared with LF. For PF, we observed little changes in the LAI and canopy openness which was attributed to the leaf retention of this evergreen species. However, a similar relation between LAI and canopy openness was found for the three forest communities: canopy openness varied inversely with LAI. The relation is exponential and significant. Therefore, canopy openness is a good indicator of LAI in forests. This result can be used to test the validity of the LAI based on remote sensing and to provide a reference for the study of the canopy heterogeneity and its effect. This also benefits modeling for fluxes of carbon, water and energy from the level of the stand to landscape. __________ Translated from Journal of Plant Ecology, 2007, 31(3): 431–436 [译自: 植物生态学报]     

Effects of bamboo stands on seed rain and seed limitation in a rainforest

Bamboos often negatively affect tree recruitment, survival, and growth, leading to arrested tree regeneration in forested habitats. Studies so far have focused on the effects of bamboos on the performance of seedlings and saplings, but the influence of bamboos on forest dynamics may start very early in the forest regeneration process by altering seed rain patterns. We tested the prediction that the density and composition of the seed rain are altered and seed limitation is higher in stands of Guadua tagoara (B or bamboo stands), a large-sized woody bamboo native from the Brazilian Atlantic Forest, compared to forest patches without bamboos (NB or non-bamboo stands). Forty 1 m² seed traps were set in B and NB stands, and the seed rain was monitored monthly for 1 year. The seed rain was not greatly altered by the presence of bamboos: rarefied seed species richness was higher for B stands, patterns of dominance and density of seeds were similar between stands, and differences in overall composition were slight. Seed limitation, however, was greater at B stands, likely as a resulted of reduced tree density. Despite such reduced density, the presence of trees growing amidst and over the bamboos seems to play a key role in keeping the seeds falling in B stands because they serve as food sources for frugivores or simply as perches for them. The loss of such trees may lead to enhanced seed limitation, contributing ultimately to the self-perpetuating bamboo disturbance cycle.     

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.     

Establishment of native tropical timber trees in monoculture and mixed-species plantations: Small-scale effects on tree performance and insect herbivory

Plantations of native timber species established on former pasture are a potential strategy to reduce the logging pressure on remnant natural forests in the tropics. Such plantations may help to mitigate or reverse the negative impacts of land degradation, and they may contribute to the long-term livelihood of livestock farmers. Planting native trees is, however, perceived as a risky activity due to limited knowledge of their performance and due to marked losses of newly established seedlings attributed to insect pests. Our study focuses on the small-scale effects of environmental heterogeneity, tree diversity and insecticide treatment on the performance of three native Central American timber species two years after establishment, and on damage inflicted by insect herbivores. Growth, survival and herbivore leaf damage were quantified for Anacardium excelsum (Anacardiaceae), Cedrela odorata (Meliaceae) and Tabebuia rosea (Bignoniaceae) planted in an experiment in Panama as (1) monocultures, (2) mixed stands, and (3) mixed stands protected by insecticides. Our study revealed that small-scale effects can have a substantial impact on the success of timber trees planted on former pasture. Growth performance and survival of the three species was strongly affected by small-scale environmental heterogeneity, which was expressed as significant differences in growth and survival among different plantation plots at the same study site. Establishment of trees in mixed stands did not have significant effects on tree survival and growth compared to pure stands, although it reduced herbivore pressure in one of the studied tree species. All tree species grew best and suffered lowest leaf damage when protected by insecticides, indicating a general influence of insect herbivory on growth of newly established trees. In contrast to growth performance, survival was not significantly affected by different management practices or herbivory. The large variability among plots in tree survival and growth, and also in the effects of management practices such as planting design and insecticide treatment, emphasizes the importance of small-scale environmental heterogeneity on tree survival and growth.     

The effects of burning and dead-wood creation on the diversity of pioneer wood-inhabiting fungi in managed boreal spruce forests

The loss of natural forest habitats due to forestry is the main reason for the decline in boreal forest biodiversity of the Nordic countries of Europe. Ecological rehabilitation may provide means to recover and sustain biodiversity. We analyzed the effects of controlled burning and dead-wood creation (DWC) on the diversity of pioneer wood-inhabiting fungi in managed Norway spruce (Picea abies) forests in southern Finland. Altogether 18 stands were first subjected to a partial cutting with ordinary logging residues in form of cut stumps and treetops left on site. The subsequent rehabilitation treatments consisted of a controlled burning applied in half of the stands and three levels of dead-wood creation (5, 30 and 60 m³ ha⁻¹). The DWC involved creation of logs; felling of whole trees to mimic downed logs formed by natural disturbance processes. Each treatment was replicated three times. Inside each stand, substrates were sampled in two different biotopes; one on mineral soil and one on mineral soil with a thin peat layer. We surveyed the fungal flora on the logs (n = 364) and the ordinary residue stumps (n = 1767) and tops (n = 845) five years after the treatments.When comparing different stands, controlled burning had a significant effect on species composition; certain species were significantly more frequent on substrates in burned stands than in unburned stands, indicating that these species were favored by controlled burning. By contrast, we found no significant effects of DWC levels or biotope on species composition or richness. When comparing different substrates, 99% of the logs hosted at least one species and the occurrence probability of certain species was significantly higher on logs than on ordinary residue stumps and tops. Yet, volume-based rarefaction analyses showed that residues were more species dense than the logs, indicating that ordinary logging residues constitute important resources for many pioneer species.We conclude that controlled burning combined with DWC have strong effects on biodiversity; it modifies the composition of the pioneer wood-inhabiting fungal species found in managed forests and may thereby also influence the further succession and diversity of the secondary fungal flora.     

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.     

Effect of species and spacing of fast-growing nurse trees on growth of an indigenous tree, Hopea odorata Roxb., in northeast Thailand

We tested the effects of species and spacing of nurse trees on the growth of Hopea odorata, a dipterocarp tree indigenous to Southeast Asia, in a two-storied forest management system in northeast Thailand. Eucalyptus camaldulensis, Acacia auriculiformis, and Senna siamea were planted as nurse trees in 1987 at spacings of 4 m × 8 m, 2 m × 8 m, 4 m × 4 m, and 2 m × 4 m in the Sakaerat Silvicultural Research Station of the Royal Forest Department, Thailand. Seedlings of H. odorata were planted in the nurse tree stands at a uniform spacing of 4 m × 4 m and in control plots (no nurse trees) in 1990. Stem numbers of some nurse trees were thinned by half in 1994. The stem diameter and height of all trees were measured annually until 1995 and again in 2007. The mean annual increment (MAI) in volume was estimated as 8.2–10.1 m³ ha⁻¹ year⁻¹ for E. camaldulensis and 0.9–1.2 m³ ha⁻¹ year⁻¹ for S. siamea, smaller than reported elsewhere. This suggests that the site properties were not suitable for them. The MAI of A. auriculiformis was 7.9–9.8 m³ ha⁻¹ year⁻¹, within the reported range. Survival rates of H. odorata in the S. siamea stands and the control plots decreased rapidly during the first 2 years but then stayed constant from 1992. In contrast, survival rates of H. odorata in the E. camaldulensis and A. auriculiformis stands were initially high (>70%), but then decreased after 1995. Stem diameter, tree height, and stand basal area of H. odorata were large in both the S. siamea stands and the control plots from then. The growth of H. odorata was largest in the 2 m × 8 m S. siamea stands. In contrast, it was restricted in the E. camaldulensis and A. auriculiformis stands owing to strong shading by their canopies. Thinning by 50% tended to facilitate the growth of H. odorata temporarily in the E. camaldulensis and A. auriculiformis stands. The stand basal areas of nurse trees and of H. odorata showed a trade-off. These results suggest that the growth of H. odorata was maximized in the S. siamea stands. We assume, however, that the growth of H. odorata could be improved even in the E. camaldulensis and A. auriculiformis stands by frequent or heavy thinning.     

Silvicultural systems for plantation mahogany in Africa: Influences of canopy shade on tree growth and pest damage

Overexploitation of the African mahoganies, Khaya anthotheca and K. ivorensis, could lead to their commercial extinction in the near future. Efforts to establish plantations to sustain supply have been discouraged by the shoot borer Hypsipyla robusta Moore. We hypothesized that there is a shade level at which Hypsipyla attack and branching are reduced, but height growth is adequate. We report on the growth of these African mahoganies and Hypsipyla attack under three different forest canopy shade levels: open (55% open sky), medium shade (26% open), and deep shade (11% open). Hypsipyla attack on K. anthotheca was 85%, 11%, and 0% attack in the open, medium, and deep shade treatments, respectively. However, growth in medium and deep shade was slow. Mean tree height for K. anthotheca was 413.8, 126.2, and 54.5 cm in the open, medium and deep shade treatments, respectively. K. ivorensis showed similar trends. The reduced growth under these shade levels limit the use of this strategy for controlling Hypsipyla attack.     

A century of riparian forest expansion following extreme disturbance: Spatio-temporal change in Populus/Salix/Tamarix forests along the Upper San Pedro River,Arizona, USA

Populus–Salix forests are a valued riparian vegetation type in western North America. These pioneer, obligate phreatophytes have declined on some rivers, raising conservation concerns and stimulating restoration plantings, but have increased on others. Understanding patterns and causes of forest change is essential for formulating conservation, restoration and management plans. Our goal was to assess spatio-temporal patterns of vegetation change on the Upper San Pedro River in semiarid Arizona, USA, one of the few undammed rivers in the region. Over 100 years ago, intense floods initiated channel incision and substantially altered hydrogeomorphology. Pioneer trees began to establish in the widening post-entrenchment zone as the surfaces began to stabilize. Using a time-series of aerial photographs (1955–2003) we quantified recent change in area of riparian cover types. Analysis indicated that wooded area in the post-entrenchment zone nearly tripled from 1955 to 2003, at the expense of bare ground, and the active channel narrowed appreciably. This forest expansion represents a long-term response to river entrenchment, with the temporal pattern influenced by recent flood cycles and biogeomorphic feedbacks. Populus–Salix have established episodically during the infrequent years with high winter flood runoff, sequentially filling available recruitment space. Older cohorts cover wide swaths of the floodplain while young trees form narrow bands lining the channel. Barring extreme flooding, the pioneer forests are expected to senesce over the coming century. An additional factor that has shaped the pattern of post-entrenchment forest expansion is anthropogenic water withdrawal. Populus–Salix forest increase has been greatest within a conservation area, where stream flows are largely perennial. In drier, agricultural sectors, Populus–Salix have declined while the more deeply-rooted Tamarix has increased. Overall, the study reveals that long-term fluctuations in pioneer forest area and age structure are common on dryland rivers, and shows how past events such as extreme floods can interact with recent environmental practices such as freshwater withdrawal to influence riparian forest patterns. This underscores the necessity of a long-term perspective for forest conservation and management.     

Epiphytic lichen diversity in late-successional Pinus sylvestris forests along local and regional forest utilization gradients in eastern boreal Fennoscandia

Pinus sylvestris-dominated forests have been heavily utilized across all of boreal Fennoscandia and the remaining natural forests are generally highly fragmented. However, there are considerable local and regional differences in the intensity and duration of past forest utilization. We studied the impact of human forest use on the diversity of epiphytic and epixylic lichens in late-successional Pinus sylvestris-dominated forests by assessing species richness and composition along both local and regional gradients in forest utilization. The effects of local logging intensity were analysed by comparing three types of stands: (i) near-natural, (ii) selectively logged (in the early 20th century) and (iii) managed stands. The effects of regional differences in duration and intensity of past forest use were analysed by comparing stands in two contrasting regions (Häme and Kuhmo–Viena). The species richness of selectively logged stands was as high as that of near-natural stands and significantly higher in these two stand categories than in managed stands. Species richness increased with the density of small understorey Picea, which correlated strongly with decreasing intensity of local forest use and increasing structural complexity of selectively logged and near-natural stands. Stands in the Häme region hosted a lower number of species, and were less likely to host many old-growth indicator species than the Kuhmo–Viena region, suggesting that species have been lost from stands in the Häme region due to a longer history of intensive forest use. We conclude that selectively logged stands, along with near-natural stands, are valuable lichen habitats particularly for species confined to old-growth structures such as coarse trees and deadwood. In landscapes where natural forests have become fragmented, the management or restoration of the remaining late-successional Pinus-dominated forests, e.g. through the use of fire, should be carefully planned to avoid adverse effects on lichen species richness.     

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.     

Long-term deer exclusion in yew-wood and oakwood habitats in southwest Ireland: Changes in ground flora and species diversity

Changes in ground flora were monitored over a 32-year period in deer exclosures in a yew-wood and a neighbouring oakwood in Killarney National Park, southwest Ireland; both woods are Annex I habitats under the European Habitats Directive. Comparison was made with unfenced plots adjacent to each of the exclosures. During the period of the study, both woods were heavily grazed by introduced sika deer (Cervus nippon). In the yew-wood exclosures, total ground flora cover increased markedly during the period of deer exclusion, the main species to increase in abundance being Rubus fruticosus agg. and Hedera helix. Herbaceous species increased initially in frequency but subsequently declined; herbaceous species had higher total cover in unfenced plots than in fenced plots at the end of the study. In the oakwood, Luzula sylvatica and Vaccinium myrtillus expanded their cover in one exclosure with cover in ferns declining, but in other exclosures where dense holly thickets developed, it was instead R. fruticosus and H. helix that became more abundant within the fences than outside. Overall, long-term fencing has caused a shift from vegetation characterised by woodland specialist to woodland generalist species and there are indications of a long-term decline in diversity. We conclude that chronic heavy grazing in these woodlands has strongly influenced the overall abundance and composition of the ground flora, but that complete exclusion of grazing is also undesirable due to potential declines in diversity of woodland specialists.     

Regeneration niches and functional traits of three common species in subtropical dune forest

South African coastal dune forests are young, highly disturbed subtropical communities where conventional models of forest dynamics may be challenged. We tested predictions from the gap-phase regeneration model by comparing seedlings of three common species representing contrasting regeneration strategies: Acacia karroo as a ruderal, Celtis africana as a coloniser of forest gaps, and Diospyros natalensis as a late-successional species. We grew seedlings under contrasting light and nitrogen levels in a greenhouse and in the field for 1 year to compare their growth and survival rates, allocation and photosynthetic traits. Species’ growth rates generally followed the expected order: Acacia > Celtis > Diospyros, but Acacia responded strongly to light and Celtis responded strongly to nitrogen, leading to cross-overs in growth rates. The plasticity of allocation and photosynthesis did not clearly differentiate the strategies, although it was greater in the light-demanding species. Acacia and Celtis tended to survive better in Acacia stands than in forest plots. Leaf-level light compensation points (LCPs) were similar for the three species in most conditions, but auxiliary data suggest Diospyros has a lower whole-plant LCP than Acacia. Growth rates and LCPs were lower than most of those reported for primary-forest species in the literature, suggesting an unusual degree of shade-tolerance in this habitat. We discuss reasons why variation in shade-tolerance may be less important here than in the prevailing model for forest regeneration and suggest other biotic factors that may help differentiate regeneration niches.