A model bridging distance-dependent and distance-independent tree models to simulate the growth of mixed forests

? It is widely believed that distance-independent tree models fail to take into account the complexity of mixed stands due to the fact that spatial structure often has a greater impact on growth and dynamics in mixed stands than in pure stands. On the other hand, distance-dependent tree models are difficult to use because they require a map of the stand, which is not only very costly but also impracticable in a routine management context.

? This paper reports the development of a model bridging distance-dependent and distanceindependent tree models, and that is designed to simulate the growth of a mixed forest. The model used distributions of the number of neighbours to reconstruct tree neighbourhoods and compute the competition indices needed as inputs to the growth model.

? Data were collected from a mixed forest of sessile oak and Scots pine in central France. The study showed that local competition indices explained a significant proportion of growth variability and that intraspecific competition was greater than interspecific competition. The model based on neighbourhood distributions gave consistent predictions compared to a distance-dependent model.

? This type of model could be used instead of distance-dependent models in management contexts.

     

Modelling site quality and individual-tree growth in pure and mixed Pinus brutia stands in north-east Greece

? Simulation tools, based on individual tree growth and mortality models can produce the most detailed predictions of forest stand development under different management schedules. These models allow the manager to predict the development of any type of stand (even- and uneven-aged, and pure and mixed stands).

? Different model approaches and predictors are required for pure even-aged or mixed uneven-aged forest stands. This study developed and compared two sets of models which enable tree-level simulation of the development of pure and mixed stands of Pinus brutia in north-east Greece. The first set of models for even-aged forestry consists of site index models, diameter growth models, tree height models, and mortality models. The second set, which is for uneven-aged forestry, uses a past growth index instead of a site index.

? The simulations and overall fitting statistics suggest that the two types of models provide realistic and accurate predictions of forest stand development and allow one to simulate the development of complex Pinus brutia stand structures in Dadia National Park forests.

? The advantages of the two approaches are discussed and it is suggested that the growth index is an effective predictor of site quality and the set of models which used such variable as predictor performed in a similar way as the models using site index, which require more information and a given stand structure (even-aged).

     

Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Climate-growth variability in Quercus ilex L. west Iberian open woodlands of different stand density

? We present the longest tree-ring chronology (141 y) of Quercus ilex L. (holm oak), and discuss the species climate-growth relationships and the influence of stand density on tree sensitivity to climate.

? Similarly to Quercus suber L., the most influential climatic variables upon holm oak growth were late spring and early summer precipitation, which enhanced growth, and high temperatures in the previous August and current July, which negatively affected growth.

? High density stands responded to similar climatic factors as low density stands, but their response was generally weaker. Holm oak sensitivity to climate has increased in recent decades, which might be related to increasing temperatures in the region. Sensitivity was higher in low density stands. Additionally, the effect of summer stress on growth seems to have increased during the same period, similarly to other species in the Iberian Peninsula, suggesting that trees are more vulnerable to climatic changes.

? Stand density could buffer the response to climate by smoothing climatic extremes. Nevertheless, the effect of competition might reverse this positive effect at the individual tree level. Precautions should be taken before providing management guidelines regarding the effect of climate change and stand density on holm oak.

     

Rank reversals in tree growth along tree size, competition and climatic gradients for four forest canopy dominant species in Central Spain

? Interspecific differences in tree growth patterns with respect to biotic and abiotic factors are key for understanding forest structure and dynamics, and predicting potential changes under climate change.

? Repeated observations from the Spanish Forest Inventory (SFI) were used to parameterize maximum likelihood estimators of tree growth as a function of tree size, competition indices and climate for Pinus pinaster, P. sylvestris, Quercus ilex and Q. pyrenaica.

? Significant responses to both biotic and abiotic factors were found, with interspecific differences in species performance along competition, temperature and precipitation gradients. Q. ilex was the species most tolerant to competition while P. pinaster was the species most sensitive to climatic variation. Species relative positions shifted along gradients of these factors with rank reversals in species performance along size, competition and climatic gradients.

? The results based on average growth matched previous forestry classifications and experimental studies on relative growth rate (RGR).

? When examining growth along studied abiotic and biotic gradients, a mismatch was found between species performance ranks as predicted by our models and information derived from previous knowledge. Those discrepancies highlight the relevance of ontogeny and environmental heterogeneity in defining species performance along competition gradients.

     

Response to canopy opening does not act as a filter to Fagus sylvatica and Acer sp. advance regeneration in a mixed temperate forest

? In mixed-species forest stands, large losses in tree species diversity often occur during the regeneration phase. In a former coppice-with-standards, we investigated whether the limiting stage in the recruitment process of advance regeneration is the immediate seedling response to canopy release. Experimental canopy gaps were opened and the survival and growth of advance seedlings (Fagus sylvatica, Acer pseudoplatanus, Acer campestre, Acer platanoides) growing in the gaps or under closed canopy were monitored for three years.

? All species responded positively and rapidly to canopy release. Survival was not affected by gap opening. Diameter increment after gap opening was similar across species, and height increment was greater for Acer platanoides and for Acer pseudoplatanus. Post-release diameter and height growth were mainly determined by pre-release seedling size. Competition from neighbouring seedlings did not affect growth in the three years following canopy opening.

? In the recruitment process of F. sylvatica and Acer sp. advance regeneration, the recovery from canopy release did not appear as a limiting step that would filter against some species. Pre-release size was the main factor accounting for post-release growth and is probably a major determinant of long-term seedling dominance.

     

A generic model of thinning and stand density effects on forest growth, mortality and net increment

? For assessing forest thinning effects at large (i.e. continental) scale, data scarcity and technical limitations prevent the application of localized or individual-based thinning models.

? Here we present a simple general framework to analyze and predict the effects of thinning on growth and mortality, including the following stand density development. The effects are modeled in relative terms so that the model can be parameterized based on any thinning experiment that includes an unthinned control, regardless of site conditions and stand age.

? The model was tested against observed thinning effects on growth and mortality from five temperate and boreal species (all species pooled r ² = 0.51). It predicted a maximum increase in net stem biomass increment of 16% and a reduction in density-related mortality of 75% compared to un-thinned conditions at stand densities of around 70% of the maximum (increment optimal density).

? A sensitivity analysis revealed overlapping ranges of near optimal density (net increment within 95% of optimal) among all tested species, suggesting that one thinning scenario can be used for many species. The simple and general formulation of thinning effects based on only five parameters allows easy integration with a wide range of generic forest growth models.

     

Disentangling the effect of size and competition: a growth model for Aucoumea klaineana

• Context Okoume (Aucoumea klaineana) is the most important timber species in Gabon, while being classified as vulnerable in the IUCN red list. Thousands of growth measurements for this species have been performed since the 1950s but, because of a lack of integrated analysis, did not bring a consistent view on its growth pattern.
• Aims This study aims at disentangling the effects of tree size and local competition on tree growth.
• Methods A growth model was fitted for okoume, using data from seven sites in Gabon and Congo. The growth model was designed to be useful for forest management, which means that it relied on variables that could be computed using forest inventory data.
• Results A lognormal growth model with a negative response to stand basal area was selected. A significant residual site effect on growth was found, with a slower growth in the sites near the border of the natural range of okoume than in the sites far from it.
• Conclusion Growth strongly responded to local stand attributes such as tree density and basal area. Growth decrease with stand age in monodominant okoume stands was correctly predicted, although tree age was not incorporated into the model.

     

Spatiotemporal patterns in seedling emergence and early growth of two oak species direct-seeded on abandoned pastureland

? Fine-scale spatial and temporal establishment patterns of direct-seeded oaks on abandoned agricultural land have been little studied despite their potential importance for long-term stand structure.

? Here we periodically monitored seedling emergence and early growth of bur oak (Quercus macrocarpa Michx.) and red oak (Q. rubra L.) on an abandoned pasture, and tested the effects of herbaceous competition, rodents, and soil physicochemical properties.

? Herbaceous competition slightly decreased diameter growth, but rodents had little impact on establishment. Red oak seedlings emerged earlier than bur oak and in a greater proportion (92% vs. 56%). Seedling emergence and early growth of both species showed significant spatial structures that were partly explained by variation in soil physicochemical properties. Bur oak was more responsive to microenvironmental heterogeneity than red oak, yet much of the variation in emergence and growth of both species remained unexplained.

? This suggests that other factors, such as acorn size or genetic variability, may exert equal or greater control than microenvironmental heterogeneity over seedling emergence and early growth of these two oak species on abandoned pastureland.

     

Litterfall production in forests located at the Pre-delta area of the Paraná River (Argentina)

? The objectives of this study were to measure litterfall production of the four floodplain forest types and to analyze the relationship between litterfall (forest type, dominant species, and organic fractions) and flood pulses.

? Litterfall production was measured in two mono-specific stands of Salix humboldtiana or Tessaria integrifolia, respectively, and two mixed forests dominated by A. inundata or Nectandra angustifolia, during 1998 and the 2000–2002 periods.

? Mono-specific stands presented similar productivities (6.8 and 6.5 Mg dry matter ha^?1 y^?1, respectively), but differed significantly from the two other. The highest litterfall production was obtained during the large flood that occurred during 1998, decreasing later throughout the study period. Leaves were the dominant fraction of litterfall, followed by branches, flowers, and fruits.

? Only S. humboldtiana, T. integrifolia and A. inundata forests showed distinct patterns of litterfall production, depending on the flooding pulse. N. angustifolia did not show a distinctive litterfall pattern.

? The forests studied here presented patterns of litter production associated with the flood pulse according to its location along a topographic gradient, that controls the litter productivity of these forests.

     

Potential use of pine plantations to restore native forests in a highly fragmented river basin

? In forests, the substitution of broadleaf species by conifers can reduce biodiversity because coniferous forests generally provide less diverse vascular understories than broadleaf forests. However, in some cases, former pine plantations might be useful for restoring native forests. We compared plant species composition on the plot scale in natural beech and mixed oak forests with that in plantations of Pinus radiata. Links between plant diversity and landscape parameters (patch size, fractal dimension and distance to the nearest patch of the same type) were investigated.

? The objective of this study was to evaluate the use of pine plantations for restoring native diversity in a zone where native forests are very fragmented.

? Similar to oak forests, plant diversity in pine plantations was high, mainly due to the presence of generalist species. Some species characteristic of oak forests also appeared in pine plantations, suggesting the onset of natural forest regeneration.

? These results suggest that pine plantations could be used to promote natural regeneration of original oak forests. Moreover, residual native stands should be conserved as important sources of native species and their seeds.

     

Species variables or environmental variables as indicators of forest biodiversity: a case study using saproxylic beetles in Maritime pine plantations

? To assess the sustainability of plantation forest management we compare two types of biodiversity indicators. We used the species richness of saproxylic beetles as a case study to test the “species” and “environmental” indicator approaches. We compared single species abundance or occurrence and deadwood volume or diversity as predictor variables.

? Beetles were sampled with flight interception traps in 40 Maritime pine plantation stands. The volume and diversity of deadwood was estimated with line intersect and plot sampling in the same stands. Predictive models of species richness were built with simple linear or Partial Least Square regressions.

? Deadwood variables appeared to be good predictors of saproxylic beetle richness at the stand-scale with at least 75% of variance explained. Deadwood diversity variables consistently provided better predictive models than volume variables. The best environmental indicator was the diversity of deadwood elements larger than 15 cm in diameter.

? By contrast, the use of “species variables” appeared to be less relevant. To reach the quality of prediction obtained with “environmental variables”, the abundance or occurrence of 6 to 7 species — some of which are difficult to identify — had to be used to build the indicator.

     

Compared effects of competition by grasses (Graminoids) and broom (Cytisus scoparius) on growth and functional traits of beech saplings (Fagus sylvatica)

? The effect of two weeds (grasses and broom, Cytisus scoparius) competition on the growth and functional traits of European beech saplings (Fagus sylvatica) was investigated in an experimental plantation in the French Massif central.

? We hypothesized that grasses would have a much more harmful effect than broom on beech growth through strong competition for soil water and nitrogen.

? A randomized block design was used with three separate blocks, each possessing three types of vegetation; grasses (mainly Agrostis capillaris, Holcus mollis and H. lanatus), broom, and bare soil as a control.

? Two years after plantation beech sapling growth was significantly greater in bare soil than with competition from grasses, and was intermediate on broom. Growth was positively correlated with beech leaf nitrogen content (N_m) and xylem water potential (Ψ_x). N and Ψ_x were dependent, with nitrogen uptake probably decreasing with increasing drought. There was no clear correlation between growth and light transmitted 10 cm below the apex of the saplings.

? Beech leaf mass on an area basis (LMA) was correlated with the three resource variables (water, nitrogen and light) and with growth, confirming that LMA is a sensitive functional trait integrating the degree of stress experienced by the saplings.

? The results suggest that broom is less competitive than grasses and, if properly managed on a sufficiently well-watered site, can generate satisfactory conditions for beech establishment and growth.

     

Site preparation and competing vegetation control affect loblolly pine long-term productivity in the southern Piedmont/Upper Coastal Plain of the United States

? A site preparation study was established in 1986 to evaluate the effect of different site preparation treatments on growth and yield of loblolly pine (Pinus taeda L.) plantations on the Piedmont and Upper Coastal Plain regions of the southern United States. Site preparation treatments included: (1) burn only, (2) chop-burn, (3) shear-pile-disk, (4) chop-herbicide-burn, (5) herbicide-burn, and (6) herbicide-burn-herbicide.

? The data from the available 19 installations at age 21 were analyzed with separate analysis of variance and a multilevel nonlinear mixed-effects modeling approach.

? The herbicide-burn-herbicide treatment significantly increased average Dbh, height, basal area and volume per hectare compared to all other treatments. The burn only treatment consistently ranked worst and was followed by the chop-burn treatment. The shear-pile-disk and chop-herbicide-burn treatments had similar overall growth pattern, and will approach the same level of pine volume as the herbicide-burn treatment.

? Loblolly pine mean annual increment in volume (m³ ha^?1 y^?1) at age 21 by treatment were: herbicide-burn-herbicide (17.9), shear-pile-disk (16.1), herbicide-burn (15.9), chop-herbicide-burn (15.4), chop-burn (14.3), and burn (11.2).

? An additional chop or herbicide treatment to the burn treatment significantly increased loblolly pine yield. Complete control of both herbaceous and woody completion enhanced long-term pine productivity.

     

Fuel and stand conditions in an isolated, unmanaged forest landscape in central Oregon

? Mature, unmanaged forests in western North America provide important reference conditions for managers, yet little is known about fuel composition and the factors that influence fuel accumulation in such stands. Our objectives were to characterize fuels in a passively managed landscape of dry forests in central Oregon and identify environmental factors influencing fuel accumulation.

? Ordination techniques and analysis of variance revealed no statistical differences in total fuel loads across a wide range of environmental conditions.

? Individual fuel size classes, however, did vary by stand location and composition. Interior stands had more 1- to 100-h fuels and snags than stands at or near the edge, stands dominated by ponderosa pine had fewer small-diameter fuels and snags, lodgepole pine stands had more 1-h fuels and snags, and white fir stands had more 10-h fuels, duff, and snag basal area.

? Tree species, density and age, and years since last fire were the most important environmental variables, explaining 23% of the variation in fuels.

? Our results present further evidence that fuel composition is highly variable at many spatial and temporal scales. They also provide useful baseline information for managers of fire-prone western forests.

     

Comparison of model forms for estimating stem taper and volume in the primary conifer species of the North American Acadian Region

? The performance of ten commonly used taper equations for predicting both stem form and volume in balsam fir [Abies balsamea (L.) Mill], red spruce[Picea rubens (Sarg.)], and white pine[Pinus strobus (L.)] in the Acadian Region of North America was investigated.

? Results show that the Kozak (2004) and Bi (2000) equations were superior to the other equations in predicting diameter inside bark for red spruce and white pine, while the Valentine and Gregoire (2001) equation performed slightly better for balsam fir.

? For stem volume, the Clark et al. (1991) equation provided the best predictions across all species when upper stem diameter measurements were available, while the Kozak (2004) and compatible taper equation of Fang et al. (2000) performed well when those measurements were unavailable.

? The incorporation of crown variables substantially improved stem volume predictions (mean absolute bias reduction of 7–15%; root mean square error reduction of 10–15%) for all three species, but had little impact on stem form predictions.

? The best taper equation reduced the predicted root mean square error by 16, 39, and 45% compared to estimates from the widely used Honer (1965) regional stem volume equations for balsam fir, red spruce, and white pine, respectively.

? When multiple taper equations exist for a certain species, the use of the geometric mean of all predictions is an attractive alternative to selecting the “best” equation.

     

Estimation of chlorophyll in Quercus leaves using a portable chlorophyll meter: effects of species and leaf age

? The potential of a chlorophyll meter (CCM-200, Opti-Sciences, Tyngsboro, MA, USA) for the estimation of total leaf chlorophyll contents was evaluated in leaves of different developmental stages for Quercus pyrenaica Willd., Q. faginea Lam., Q. suber L. and Q. ilex L. subsp. ballota (Desf.) Samp.

? For all species and leaf stages, a strong non-linear relationship (P < 0.0001) was found between the chlorophyll content index (CCI) and leaf chlorophyll content per unit area. In all species, the slope of the CCI-chlorophyll content relationship varied during leaf development, suggesting that caution should be exercised when using the CCM-200, since the interpretation of CCI readings should be limited to samples of similar leaf age. Also, the data indicate that the models should also be speciesspecific, owing to differences in the intercept and/or slope of the equations.

? The addition of fresh LMA as a complementary parameter improved the accuracy of chlorophyll estimation in non-species-specific equations to a considerable extent when leaf samples of similar development were considered.

? After due consideration, it may be concluded that the CCM-200 chlorophyll meter is a reliable method for acquiring an estimation of chlorophyll contents in Mediterranean Quercus species, with potential use as an effective tool in forest management.

     

Which factors most influence heartwood distribution and radial growth in plantation teak?

? To investigate the effect of climate on radial growth in young plantation grown teak (Tectona grandis L.), growth ring width was measured in 105 trees and correlated to precipitation and temperature data.

? The social status of trees within the stand was also determined and cross-sectional area (CSA) for the trunk correlated to the proportion of heartwood (HW) within the tree. HW develops asymmetrically in leaning stems of some conifer species, but it is not known if this phenomenon also occurs in broadleaf species. Therefore, we measured HW proportion in leaning and straight stems, along with the number of growth rings in the HW.

? Annual ring width depended strongly on mean monthly temperature during the rainy season and the most significant relationships were found corresponding to the months of June and July. With regard to the weaker relationship between precipitation and radial growth, correlations were highest during the period of bud-break at the beginning of the rainy season.

? The very high stand density affected radial growth, particularly in suppressed trees, which responded little to thinning operations. HW formation was greatest in dominant trees, and was highly regressed with stem CSA.

? Therefore, rapid growth of young stands should be encouraged by reducing stand density. Asymmetric HW formation occurred in both leaning and straight trees, and was significantly greater along the upper sides of leaning stems. It is probable that this eccentric HW formation is linked to mechanical loading on the tree.

     

Ecology of Armillaria species on silver fir (Abies alba) in the Spanish Pyrenees

? We describe the distribution and the ecology of three Armillaria species observed in silver fir (Abies alba) forests of the Pyrenees.

? We surveyed the presence and abundance of Armillaria above and belowground in 29 stands. Isolates were identified by the PCR-RFLP pattern of the IGS-1 region of their ribosomal DNA. We measured several ecological and management parameters of each stand in order to describe Armillaria infected sites.

? Armillaria cepistipes was the most abundant of three species observed. Armillaria gallica was dominant in soils with a higher pH and at lower elevations. Armillaria ostoyae seemed to be more frequent in stands where A. alba recently increased its dominance relative to other forest tree species. Thinning activities correlated with an increased abundance of Armillaria belowground. In 83% of the stands the same Armillaria species was observed above and belowground.

? It seems that in a conifer forest, A. cepistipes can be more frequent than A. ostoyae, a virulent conifer pathogen. Since logging is related to a higher abundance of Armillaria in the soil, the particular Armillaria species present in a given stand could be considered an additional site factor when making management decisions.

     

Development of alder carr after the abandonment of wet grasslands during the last 70 years

? The secondary succession of wet grasslands to communities of alder carr dominated by Alnus glutinosa was recorded in different parts of Europe during the 20th century. However, knowledge of such development of alder carr remains insufficient.

? The development of alder carr was reconstructed at five sites in the Czech Republic, using historical aerial photographs and methods of dendrochronology. The aims were to investigate the succession from wet grasslands to alder carr at sites previously intensively managed for agriculture and to find out the role of fluctuations in the groundwater table, caused by artificial drainage channels, in the observed stand dynamics and tree growth.

? The spread of forest (i.e., an increase in forest cover) predominated until the 1970s at all sites. This trend was disrupted by a large-scale dieback of forest stands in four of the five sites after the 1970s, followed by an increase in patch heterogeneity, as indicated by landscape metrics. The radial growth increment in Alnus glutinosa has been affected predominately by local environmental factors, probably including the changing degree of waterlogging. Forest dieback was presumably connected with a lesser extent of drainage channels.

? Our results indicate that observed successional pathways at sites of alder carr were probably caused by local changes in the groundwater table.