Impact of spatial heterogeneity of neighborhoods on long-term population dynamics of sugar maple (Acer saccharum)

We evaluated the impact of spatial heterogeneity resulting from disturbance and neighbor density on long-term population dynamics of sugar maple (Acer saccharum) in Brownfield Woods, an old-growth forest remnant in central Illinois. All trees with diameter at breast height (DBH) ≥ 7.6 cm in eight discrete quadrats (48 m × 68 m) and one large quadrat (216 m × 260 m) were mapped in Brownfield Woods in 1951, 1988, and 2001. Two sets of neighbor-specific transition matrices were constructed for two census periods (1951–1988 and 1988–2001) based upon historical maps of trees. Different neighborhoods defined by the occurrence of disturbance and neighbor density resulted in major differences in subsequent demography and population dynamics of A. saccharum in Brownfield Woods during 1951–2001. Disturbance subpopulations with high neighbor density demonstrated greater population growth than the other subpopulations and were characterized by relatively high recruitment and mortality. Dutch elm disease subpopulation shared similar characteristics. In contrast, non-disturbance subpopulations demonstrated census-specific demography. Using matrix modeling, we demonstrated the importance of spatial heterogeneity at the scale of neighboring trees to the dynamics at the scale of tree populations.     

Canopy disturbance and tree recruitment over two centuries in a managed longleaf pine landscape

Disturbance history was reconstructed across an 11300 ha managed longleaf pine (Pinus palustris Mill.) landscape in southwestern Georgia, USA. Our specific objectives were to: (i) determine forest age structure; (ii) reconstruct disturbance history through the relationship between canopy disturbance, tree recruitment and growth; and (iii) explore the relationship between canopy disturbance and climate. Age structure, canopy disturbance events and initial growth patterns at coring height were examined by randomly sampling 1260 trees in 70 1.3 ha plots. Principal component analysis was used to group plots with similar age structures to gain insight into the dynamics between canopy disturbance and recruitment. Disturbance events were detected by large and rapid increases in radial growth. We tested the following hypothesis to investigate whether these growth increases could have been triggered by improved climatic conditions: precipitation and drought are positively correlated to radial growth releases. Only four stands (comprising <6% of the study area) had an even-aged structure. Further, tree recruitment prior to European settlement indicates that longleaf pine naturally recruited into areas 1.3 ha or less, supporting early-20th century observations that the primary longleaf pine forest was uneven-aged. Contrary to our hypothesis, growing season precipitation and drought was significantly and negatively correlated with canopy disturbance (radial growth releases), which indicates that a reconstruction of disturbance history could proceed with some confidence. Most trees sampled were recruited at coring height from 1910 to 1935. Of the 67 canopy disturbances detected from 1910 to 1935, the average growth release ranged from 139 to 277% per half decade suggesting the occurrence of large canopy disturbances. Rapid initial growth patterns of young trees during these years show evidence of reduced overstory competition and support the detected disturbance intensity. Our reconstruction of stand dynamics is markedly similar to independent records of local oral and written history, which gives an additional set of evidence that the disturbance detection methodology used can be useful in open-canopied forests. Stands with multiple cohorts reveal a mix of continuous minor and major canopy disturbances leading to continual tree recruitment, suggesting their applicability as models for long-term forest management. The significant relationship between climate and disturbance in our data suggests that with the expected warming over the next 100 years, climatic impacts on stand dynamics should be incorporated into long-term longleaf pine forest restoration and management.     

Enrichment planting as a silvicultural option in the eastern Amazon: Case study of Fazenda Cauaxi

Liana-dominated forest patches constitute 15–20% of old-growth forests in the Eastern Amazon but are generally excluded from management for timber production. Here we ask if liana-dominated patches may be brought into production by clearing lianas and conducting enrichment planting (EP) of native timber species. We present growth results from 8 years of such EP trials. Rapid growth and low mortality of all species in this study suggest that EP in cleared liana patches can contribute to timber stocks in second and third harvests of managed forests. The most vigorous individuals of Parkiagigantocarpa and Schizolobium amazonicum in each enrichment site grew more than 1 cm diameter per year (rates were initially >2 cm yr⁻¹), and attained dominant canopy positions and diameters equal to those of small canopy trees in the surrounding forest within 8 years of planting (mean dbh ∼18 cm and ∼20 cm, respectively, at year 8). Limited data on Ceiba pentandra plantings indicate a similar trajectory for this species (dbh ∼40 cm in 8 years). The most vigorous Swietenia macrophylla grew at least 1 cm per year in enrichment plots (mean dbh ∼10 cm in 8 years), but take longer to attain dominant positions. Tabebuia serratifolia may take much longer to reach the canopy than other species tested (rates <1 m yr⁻¹). We attribute the excellent performance to light availability; planting in intact soil with minimal compaction and abundant organic material; and low competition rates maintained by periodic thinning of competing vegetation.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.     

Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

Patterns of spread of Pinus contorta Dougl. ex Loud. invasion in a Natural Reserve in southern South America

Pinus species, which have formed the foundation of commercial forestry industry in many countries, are known to be invasive in natural ecosystems, especially in the Southern Hemisphere. Pinus contorta is considered one of the most aggressively invasive plantation species. In this paper we aim to: (a) determine patterns of P. contorta invasion in relation to its size and age structure and spatial distribution and (b) determine the effect of vegetation cover on its regeneration. For this purpose, we have chosen the Malalcahuello National Reserve in south-central Chile, which is dominated by Araucaria araucana forests. In the three P. contorta trial plots, attributes were measured in order to describe its current structure. We selected one of the three P. contorta trial plots and set eight 50 m wide and 125 m centrifugal transects starting at the North Azimuth (0°) orientation and then one each 45°. In each transect 25 circular plots of 2.5 m radius were established every 25 m. In each plot, we measured collar diameter (CD) for each P. contorta individual and adjusted a diameter–age function. We recorded the presence of cones for each individual P. contorta within the plots and the total number of P. contorta individuals. In each plot, we measured percent vegetation cover (grass, understory and canopy). The selected parent stand showed a decrease in density from the original plantation spacing, from 2500 to 150 plants ha⁻¹. Of all sampled individuals, only a 38% had cones. We found reproductive structures in trees as young as 5-year-old. Regeneration was found in all transects. Mean density for the area was 1600 plants ha⁻¹, and the greater plant number was found in the West transect, reaching an average of 6600 plants ha⁻¹. Through the interpolation performed with the kriging method, a map of the area with the spatial gradient of plant density was obtained. Naturally regenerated individuals of P. contorta occupied an area of 78 ha where the most distant individual is located in the southeast direction at an average distance of 1200 m from the three study plots. We determined that a positive association exists between P. contorta and the species A. araucana and Nothofagus antarctica. The capacity for early and consistent reproduction and the establishment of individuals dispersed at great distances from the original plots allow us to conclude that P. contorta has great potential as an invasive species in forests of this area of the Andes of southern South America. Invasion of P. contorta has many important implications for the conservation of native forests in our region including diminished regeneration of A. araucana.     

Winter litter disturbance facilitates the spread of the nonnative invasive grass Microstegium vimineum (Trin.) A. Camus

We investigated the impacts of winter litter disturbance on the spread of the nonnative invasive plant Microstegium vimineum (Trin.) A. Camus through experimental removals. We hypothesized that light penetration through the litter layer facilitates the spread of M. vimineum in forested systems. Our objective, therefore, was to quantify M. vimineum spread following litter removal. Linear spread and cover expansion from established M. vimineum patches was documented for one growing season under intact, undisturbed hardwood canopies within plots receiving one of two treatments. Treatments included litter removal (hereafter “removal”) and no litter removal (hereafter “undisturbed”). After one growing season, plots receiving the removal treatment experienced a spread of M. vimineum 4.5 times greater than plots receiving the undisturbed treatment (P < 0.0001; 1.66, and 0.37 m expansion, respectively). Cover expansion (measured as percent cover in 0.5 m² blocks at 0.5, 1, 1.5, and 2 m from established M. vimineum) averaged 16, 4, 0, and 0%, respectively, for the undisturbed treatment and 87, 64, 31, and 9%, respectively, for the removal treatment. Differences existed in cover expansion between treatments at the 0.5, 1, and 1.5 m distances (P < 0.0001, P < 0.001, and P = 0.01, respectively). Our results suggest that winter litter removal as a result of harvest activities, floodwater scour, or animal activities can drastically increase M. vimineum spread and may enhance potential ecological impacts of invasions by increasing M. vimineum percent cover. Previous studies have shown that M. vimineum responds to canopy removal with dramatic increases in biomass. This study suggests one mechanism facilitating rapid expansion of M. vimineum following site disturbance, and indicates that M. vimineum can experience rapid growth in response to site disturbance even in the absence of canopy removal.     

Canopy gap dynamics and development patterns in secondary Quercus stands on the Cumberland Plateau, Alabama, USA

Forest disturbances of various spatial extents and magnitudes shape species composition, structure, and stand development patterns. The disturbance regimes of most complex stage hardwood stands of the deciduous forests of eastern North America are typified by asynchronous and localized disturbance events. The overwhelming majority of gap-scale disturbance studies in hardwood forests of the region have analyzed late-successional stands. As such, there is a paucity of data on gap dynamics in hardwood stands prior to a complex developmental stage. We quantified biophysical characteristics of 60 canopy gaps in secondary Quercus stands on the Cumberland Plateau in Alabama to analyze gap-scale disturbance processes in developing systems. We found most gaps (90%) were caused by the removal of a single tree. Of the three gap formation mechanisms, snag-formed gaps were most common (40%). However, based on the number of uprooted and snapped stems we speculate that wind was also an important disturbance agent in these stands. Gap size and shape patterns were similar to what has been reported in other hardwood forests of the southern Appalachian Highlands. We did not find differences in gap size or shape based on formation mechanisms; a finding that may be related to the number of single-tree gap events. Gaps projected to close via subcanopy recruitment were significantly larger than those projected to close through lateral crown expansion. Most gaps (65%) were projected to close by lateral crown expansion of gap perimeter trees. However, the number of gaps projected to fill by subcanopy recruitment indicated the stands were approaching a transition in their developmental stage. Gap-scale processes modify residual tree architecture and stand structure. Through time these alterations result in progressively larger gaps, eventually reaching a size when most will fill by subcanopy recruitment, thus marking the complex stage of development. Gap capture by Quercus was restricted to relatively xeric sites that did not contain abundant shade-tolerant mesophytes in the understory. However, the majority of gaps contained abundant subcanopy Fagusgrandifolia, Acer saccharum, and Acer rubrum leading us to project that the forest will undergo a drastic composition shift under the current disturbance regime. Liriodendron tulipifera was projected to capture several relatively small gaps illustrating the role of topography on gap closure mechanisms.     

The effects of sudden oak death on foliar moisture content and crown fire potential in tanoak

The introduction of non-native pathogens can have profound effects on forest ecosystems resulting in loss of species, changes in species composition, and altered fuel structure. The introduction of Phytophthora ramorum, the pathogen recognized as causing Sudden Oak Death (SOD), leads to rapid decline and mortality of tanoak (Lithocarpus densiflorus) in forests of coastal California, USA. We tracked foliar moisture content (FMC) of uninfected tanoaks, SOD-infected tanoaks, SOD-killed (dead) tanoaks, and surface litter for 12 months. We found that FMC values differed significantly among the three categories of infection. FMC of uninfected tanoaks averaged 82.3% for the year whereas FMC of infected tanoaks had a lower average of 77.8% (ANOVA, P = 0.04). Dead trees had a significantly lower FMC, averaging 12.3% (ANOVA, P < 0.01) for the year. During fire season (June–September), dead tanoak FMC reached a low of 5.8%, with no significant difference between dead canopy fuels and surface litter (ANOVA, P = 0.44). Application of low FMC values to a crown ignition model results in extremely high canopy base height values to escape crown ignition. Remote estimation of dead FMC using 10-h timelag fuel moisture shows a strong correlation between remote automated weather station (RAWS) 10-h timelag fuel moisture data and the FMC of dead leaves (R² = 0.78, P < 0.01). Results from this study will help refine the decision support tools for fire managers in SOD-affected areas as well as conditions in other forests where diseases and insect epidemics have altered forest canopy fuels.     

Interannual variation in competitive interactions from natural and anthropogenic disturbances in a temperate forest tree species: Implications for ecological interpretation

Competition is a major determinant of plant growth and is often used in studies of tree growth and species coexistence. However, these approaches are usually temporally static, i.e., assessed at a single point or period in time. While constantly changing forest conditions due to natural and human-induced disturbances potentially alter competition among individuals, static approaches cannot qualify the temporal variability of competitive interactions. Here we present a longitudinal analysis of competitive interactions among trees and discuss the implication of our results for ecological interpretation.Spatially-explicit tree growth data were obtained from 18 study plots (0.4 ha each) in sugar maple (Acer saccharum Marsh.) stands in Quebec, Canada. During the studied period (1980-2003), these stands had been disturbed by insect outbreaks (forest tent caterpillar, Malacosoma disstria Hubner) and by commercial partial harvest. We analyzed radial growth rates (outcome of competition) on an annual basis and as a function of tree biology (bole diameter, crown position), competition (above- and belowground competition from neighbours) and environmental conditions (light availability, harvest disturbance).Competitive interactions changed throughout the studied period. Canopy disturbance from partial harvest interacted with defoliators and influenced competition symmetry by favoring smaller trees.Competitive interactions seemed to have switched from below- to above-ground following canopy recovery after harvest. Release from competition due to partial harvest increase neighbourhood size (radius of effective competition) and enhanced the competitive pressure from larger individuals.The temporal variability in parameter estimates may be used for setting confidence intervals on competitive success (growth rates), thereby yielding a more robust basis for ecological interpretation. Our results also show that temporal variability in competitive interactions could contribute to the maintenance of high tree species diversity and structural complexity in some ecosystems by temporally altering species-specific responses to environmental change and disturbance.     

Spatial associations among tree species in a temperate forest community in North-western Spain

Analyzing spatial patterns in plant communities may provide insights in the importance of different processes for community assembly and dynamics. We applied techniques of spatial point pattern analysis to data from a fully mapped plot of a temperate forest community (Corylus avellana, Crataegus monogyna, Fagus sylvatica, Ilex aquifolium and Taxus baccata) in North-western Spain to conduct a community wide assessment of the type and frequency of intra and interspecific spatial association patterns. We first explored the overall intra and interspecific patterning, and then classified the types of association patterns at various neighbourhoods. By conditioning on the larger scale pattern we then explored small-scale (0–15 m) intraspecific and interspecific patterns. Association patterns varied from strong positive association at small scales to, as a by-product, repulsion at intermediate scales. Surprisingly, there were no negative associations at small scales, but trees were arranged in multi-species clumps, up to 2.5 m in diameter and comprising a few individuals. Ilex and Corylus, the understorey species, were frequently involved in the clumps, showing positive small-scale association with the other tree species. Our analyses highlighted that animal mediated seed dispersal, interspecific facilitation and perturbation processes may operate successively to shape tree distributional patterns, although their relative importance vary among species. Given the complexity of the patterns described and the current threats to some of the species studied, directed experiments in the field are needed to further elucidate some of the hypotheses derived.     

Long-term dynamics of a mixed conifer stand in Slovenia managed with a farmer selection system

The single-tree selection system is an important option for management of Norway spruce (Picea abies (L.) Karst.) and silver fir (Abies alba Mill.) forests because it provides continuous cover, requires low investments for tending, and promotes natural regeneration as well as high stand resistance and elasticity. It is often regarded as a very conservative system that usually results in only minor spatiotemporal changes in forest structure and composition. We studied management history, structural changes, regeneration dynamics, and light climate of a traditional single-tree farmer selection silver fir-Norway spruce forest (site typology Bazzanio-Abietetum). Stand structure was analyzed on five 0.25 ha permanent plots in 1994, 2001, and 2008. Regeneration density and height growth, forest floor vegetation, and light climate were also assessed on 1.5 × 1.5 m regeneration subplots in 2001 and 2008. Tree cores extracted from dominant trees from both species in two plots were used for reconstructing stand history and age structure of the canopy layer. We documented the forest response to three types of selection management regimes: excessive, normal, and conservative. Excessive management with harvest intensity significantly above the increment was documented until the late 1950s, including two peaks of heavy fellings (diameter limit cut) in the 1880s and 1930s, which favoured establishment of Norway spruce and released regeneration. The period that followed was characterized by normal selection management, but was nevertheless marked by a decline of silver fir as a result of air pollution and several droughts. This led to sanitary fellings that were carried out from the late 1970s to the early 1990s. In the last two decades conservative management followed, which led to suppression and decline of regeneration, especially of Norway spruce, and loss of selection structure. Although we recorded lower regeneration potential of silver fir compared with Norway spruce within the seedling category, silver fir outcompeted Norway spruce within the small-sized tree category (1 cm < dbh ? 10 cm) because of its superior height growth in low light levels (diffuse light <6%) and occupied a greater share of the canopy. Nevertheless, we anticipate that over the long-term the low light regime will also cause regeneration decline of silver fir and broadleaves. Our research revealed significant structural changes in a single-tree farmer selection forest during the last 150 years. These were a result of variable management regime and environment. A farmer single-tree selection system could better mimic the natural disturbance regime if spatiotemporal combinations of diverse felling regimes would be used.     

Tree biomass estimation in regenerating areas of tropical dry vegetation in northeast Brazil

Allometric equations have been developed for various different vegetation types but have rarely been validated in the field and never for dry tropical forest such as caatinga. In three areas of semi-arid Brazil, with regenerating caatinga vegetation, we measured and weighed twelve hundred individuals of four tree species and used the data to validate equations previously determined in mature caatinga. They and several other equations developed for tropical vegetations overestimate the biomass (B) of trees from the regeneration areas by more than 20%, possibly because these trees have reduced crowns, with lower branch masses. We then determined new allometric equations for them, validating equations for one site against data of the others and pooling the data if they were cross-validated. The best equations were power ones, based on diameter at breast height (D), with little improvement by including height, crown area and/or wood density (Caesalpinia pyramidalis, B = 0.3129D^1.8838; Croton sonderianus, B = 0.4171D^1.5601; Mimosa ophthalmocentra, B = 0.4369D^1.8493; and Mimosa tenuiflora, B = 0.3344D^1.9648 and 0.4138D^1.7718).     

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

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

Water use by Tabor and Kermes oaks growing in their respective habitats in the Lower Galilee region of Israel

We estimated water use by the two main oak species of the Lower Galilee region of Israel—Tabor (Quercus ithaburensis) and Kermes (Quercus calliprinos)—to develop management options for climate-change scenarios. The trees were studied in their typical phytosociological associations on different bedrock formations at two sites with the same climatic conditions. Using the heat-pulse method, sap flow velocity was measured in eight trunks (trees) of each species during a number of periods in 2001, 2002 and 2003. Hourly sap flux was integrated to daily transpiration per tree and up-scaled to transpiration at the forest canopy level. The annual courses of daytime transpiration rate were estimated using fitted functions, and annual totals were calculated. Sap flow velocity was higher in Tabor than in Kermes oak, and it was highest in the youngest xylem, declining with depth into the older xylem. Average daytime transpiration rate was 67.9 ± 4.9 l tree⁻¹ d⁻¹, or 0.95 ± 0.07 mm d⁻¹, for Tabor oak, and 22.0 ± 1.7 l tree⁻¹d⁻¹, or 0.73 ± 0.05 mm d⁻¹, for Kermes oak. Differences between the two oak species in their forest canopy transpiration rates occurred mainly between the end of April and the beginning of October. Annual daytime transpiration was estimated to be 244 mm year⁻¹ for Tabor oak and 213 mm year⁻¹ for Kermes oak. Adding nocturnal water fluxes, estimated to be 20% of the daytime transpiration, resulted in total annual transpiration of 293 and 256 mm year⁻¹ by Tabor and Kermes oaks, respectively. These amounts constituted 51% and 44%, respectively, of the 578 mm year⁻¹ average annual rainfall in the region. The two species differed in their root morphology. Tabor oak roots did not penetrate the bedrock but were concentrated along the soil–rock interface within soil pockets. In contrast, the root system of Kermes oak grew deeper via fissures and crevices in the bedrock system and achieved direct contact with the deeper bedrock layers. Despite differences between the two sites in soil–bedrock lithological properties, and differences in the woody structure, annual water use by the two forest types was fairly similar. Because stocking density of the Tabor oak forests is strongly related to bedrock characteristics, thinning as a management tool will not change partitioning of the rainfall between different soil pockets, and hence soil water availability to the trees. In contrast, thinning of Kermes oak forests is expected to raise water availability to the remaining trees.     

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.     

Survival and early growth of mixed forest stands installed in a Mediterranean Region: Effects of site preparation intensity

In Mediterranean environments, availability of water and nutrients are the main factors limiting the success of afforestation. As part of a wider project, an experiment was established in Northeast Portugal, aiming at testing the effect of several site preparation techniques on plant survival and growth (height and diameter) in a newly installed mixed forest stand. Results presented regard plant response during 42 months after plantation. The experimental protocol consisted in seven treatments described by mechanical operations that rank soil disturbance intensity from none to high, set in plots of 375 m², randomly distributed in three blocks, in different topographic positions (gentle slope plateau, moderate slope shoulder, and steep mid-slope). Pseudotsuga menziesii (PM) and Castanea sativa (CS) forest species were planted in a 4 m × 2 m scheme and in alternate rows with 12 plants on each row per plot, summing up 72 plant per specie and treatment at start of the experiment. The results show that: (i) the highest mortality was observed immediately after the plantation and before the dry season, on the lowest intensity treatments; (ii) after the dry season, the highest mortality was also observed in treatments with the lowest intensity of soil disturbance, while the lowest values were found on the intermediate intensity treatments; (iii) during the experimental period, the effect of treatments on plant growth (height and diameter) was statistically significant; however, experimental results do not lead yet to a clear quantitative relationship between soil disturbance intensity due to site preparation and plant response under the conditions tested.     

Recruitment limitation in forests: Lessons from an unprecedented mountain pine beetle epidemic

Since the mid-1990s the forests of central British Columbia have undergone an unprecedented Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) (MPB) epidemic that has resulted in extensive mortality of canopy lodgepole pine (Pinus contorta var. latifolia Engelm.). This study investigated how seed-source availability, seedbed substrate, overstory structure, and time since MPB attack interact to affect post-MPB seedling recruitment of the dominant tree species of these forests. In addition to post-MPB recruitment, these forests may be regenerated by trees established in the understory prior to MPB disturbance. Accordingly, we examined abundance and patterns of all regeneration less than 130 cm tall. We found post-MPB recruitment was sparse. Subalpine fir (Abies laciocarpa (Hook.) Nutt.) comprised the majority of the post-MPB recruitment. It increased with local parent tree basal area and increased strongly with proximity to a major seed source. This resulted in a patchy distribution for subalpine fir post-MPB regeneration. Lodgepole pine post-MPB recruitment was limited by overstory shading. Recruitment of pine decreased as the total overstory basal area increased. Interior spruce (Picea glauca × engelmannii) post-MPB recruitment was similarly limited by total overstory basal area. Seedbed substrates were uniform and dominated by moss. Substrate type distribution did not change as time since MPB disturbance increased. The overall low post-MPB recruitment observed was likely due to a lack of disturbance to the moss-dominated forest floor. Moss is known to be a poor substrate in northern forests. The distribution of all regeneration less than 130 cm tall showed the same trends as the post-MPB regeneration. We believe the post-MPB seedling recruitment dynamics of these forests was not substantially changed from conditions prior to MPB disturbance. There was no pulse of regeneration up to 10 years post-MPB disturbance. Unless this changes, future stand structure will be dominated by the seedling bank established prior to the MPB epidemic. Subalpine fir dominated the seedling bank (68%) and post-MPB recruitment (94%). This suggests that MPB-disturbed forests are undergoing a substantial shift in landscape-level species composition.     

Fragmentation and spatial genetic structure in Tabebuia ochracea (Bignoniaceae) a seasonally dry Neotropical tree

In this study we investigate the effect of fragmentation and disturbance on the spatial genetic structure, heterozygosity and inbreeding in Tabebuia ochracea (Bignoniaceae) in a seasonally Neotropical dry forest in the medium São Francisco River basin, Centre-East Brazil, based on the polymorphism at seven microsatellite loci. Four populations with different histories of disturbance and fragmentation were sampled: two continuous population (CP1 and CP2), with no history of recent disturbance and two fragmented and isolated population (FP1 and FP2), with recent history of disturbance due to logging for pasture establishment. Fragmented and continuous populations did not differ in any estimated parameter. However, all populations showed low levels of polymorphism and genetic diversity and high levels of inbreeding. Also, no spatial genetic structure was detected for populations using SPAGeDI software and no differentiation between these four populations was detected by Bayesian analyses performed with STRUCTURE software (K = 1). Differentiation measure by Wright's θ (0.032) and Hedrick G_ST (0.032) were significant but low. Our results strongly suggest that continuous populations are seed sources for the fragmented populations and that fragmentation and disturbance have been affecting these populations of T. ochracea in the Centre-East Brazil, leading to low levels of polymorphism and genetic diversity, and high inbreeding. Therefore, conservation efforts should increase in this region, with a reduction of agriculture expansion and the remove of cultivated areas and cattle from the Mata Seca and Lagoa do Cajueiro State Parks.