Modelling cork oak production in Portugal

The layout of 32 permanent plots in the most important cork production regions of Portugal and the measurement of the most representative tree parameters were performed for statistically sound cork weight modelling. The establishment of cork weight equations is presented for the different cork oak study areas considered alone or grouped according to the most representative tree parameters. For the presented cork weight models was chosen independent variable the total stripping surface (artmax). For the selection criteria five statistic were used, including PRESS, APRESS and Mallows's statistic. The crossvalidation of the best classified models was performed and to discuss the importance of the “study area” factor, a regression analysis with “dummy” variables was performed.     

Analysis of variables influencing tree cork caliper in two consecutive cork extractions using cork growth index modelling

The influence of tree and stand variables, debarking intensity, and precipitation on the caliper of cork produced by a tree and on the evolution of cork caliper between consecutive cork extractions was researched. A total of 370 cork samples were collected in 23 permanent plots distributed across the cork production area in Portugal, covering a period from 1984 to 2010. Cork growth was evaluated using the cork growth index (cgi), defined as the radial width of the first eight complete years of cork growth after stripping. The differences in mean cgi at plot level between two consecutive cork growth periods (cgp) were assessed using nonparametric tests. A mixed model approach was used due to the nested structure of the data for modelling cgi value at tree level. The cgi of two consecutive cork extractions is linearly related at tree level. At plot level, the mean value of cgi decreased in 5 out of the 23 plots and increased in 2 plots for α = 0.05. The number of debarked branches and the variation of precipitation between two cgp were the variables that explained the largest part of the cgi evolution. However, significance of plot random parameters indicates that other variables are involved in the tree cgi evolution, pointing out to the need of further research. Tree size and precipitation during the cgp are related to the individual tree cgi. The effect of increasing stand density and debarking intensity on cork growth was not clear. Long term research based on permanent plot measurements and research trials is needed to clarify the impact of tree competition and debarking intensity.     

Mediterranean cork oak wooded grasslands: synergies and trade-offs between plant diversity,pasture production and soil carbon

Mediterranean wooded grasslands that emerge from silvopastoral activities are multifunctional systems that result in high biodiversity and offer ecosystem services such as forage production and soil carbon sequestration. During 3 years, ten grazed wooded grassland fields were studied in the Berchidda–Monti long-term observatory, located in NE Sardinia, Italy, with the aim of exploring the synergies and trade-offs between biodiversity and selected ecosystem services. Positions below and outside the canopy of three cork oak trees in each field were randomly selected to compare seasonal pasture production, pasture utilization rate by animals, botanical composition, biodiversity indicators (Shannon index and plant species richness) and soil organic carbon. In autumn, dry matter production of pasture was similar in the two positions; in two winters out of three it was greater below the trees than outside, and in spring it was greater outside than below the trees. While plant species richness and Shannon index were not significantly influenced by the position, the overall wooded grassland plant species richness was 31% higher than that outside of the tree crown. The soil organic carbon content in the 0–40-cm soil layer was also higher below the trees. Our findings highlight that if the main purpose of the wooded grasslands is to provide forage for grazing animals rather than conserving and/or enhancing plant diversity and soil fertility, the presence of trees constrains the overall forage productivity, although the greater forage availability in winter under the trees can contribute to improve the seasonal distribution of forage production.     

Variables influencing cork thickness in Spanish cork oak forests: A modelling approach

In this study, we evaluate the influence of different variables on cork thickness in cork oak forests. For this purpose, first we fitted a multilevel linear mixed model for predicting average cork thickness, and then identified the explanatory covariates by studying their possible correlation with random effects. The model for predicting average cork thickness is described as a stochastic process, where a fixed, deterministic model, explains the mean value, while unexplained residual variability is described and modelled by including random parameters acting at plot, tree, plot × cork harvest and residual within-tree levels, considering the spatial covariance structure between trees within the same plot. Calibration is carried out by using the best linear unbiased predictor (BLUP) theory. Different alternatives were tested to determine the optimum subsample size which was found to be appropriate at four trees. Finally, the model was applied and its performance in the estimation of cork production was tested and compared with the cork weight model traditionally used in Spain.     

Analysis of spatial patterns of oak decline in cork oak woodlands in Mediterranean conditions

? Cork oak mortality is a recurrent problem in southwestern Portugal. Despite the perception of increasingly visible damage in oak woodlands on drought-prone sites, the role of the various environmental factors in their decline is not clear.

? To describe the spatial patterns of cork oak (Quercus suber L.) mortality, a cork oak mortality index (MI) was determined for each landscape feature (agroforestry system, soil type, slope and aspect) using a GIS approach. To achieve this goal, a logistic regression model was formulated analyzing interactions between landscape attributes and allowing a prediction of cork oak mortality.

? Maximum values of MI were found in (i) shrublands and open woodlands with shrub encroachment (MI 6 and 3, respectively), where competition for soil water between tree and understory increases; and (ii) on lower slopes in the rounded hilltops and smooth hillsides or shallow soils where access to groundwater resources during summer drought is difficult.

? The model highlighted the importance of the agroforestry systems on cork oak mortality and may be used to identify sensitive areas where mitigation actions should be employed in a scenario of increasing drought severity in these Mediterranean ecosystems.

     

Physiological responses of cork oak and holm oak to infection by fungal pathogens involved in oak decline

The aim of this research was to study the changes in net photosynthesis and stomatal conductance values in 3‐year‐old cork oak and holm oak seedlings growing in natural conditions and inoculated with Apiognomonia quercina, Biscogniauxia mediterranea, Botryosphaeria corticola and Pleurophoma cava. Throughout the 4‐month experimental period, the evolution of visual external symptoms and the values of physiological variables were periodically recorded. All pathogens caused stem lesions around the infection point; however, the lesions caused by B. corticola were longer in both oak species. On cork oak seedlings, all pathogens induced a significant and gradual reduction in net photosynthesis and stomatal conductance values, whereas other physiological disturbances were induced only by B. corticola infections on holm oak seedlings.     

The effect of cork removal on the radial growth and phenology of young cork oak trees

The effect of the removal of cork was studied in 11-years old cork oak trees (Quercus suber L.), growing in favorable conditions, in relation to phenology and radial growth during two years. Longevity of leaves was 14–15 months (1996, 1998) and 10 months (1997). Bud burst started in mid-February and leaf flushing in April, extending until June. Neither a distinct two-period flushing nor an autumn bud burst or leaf flushing were observed. Radial growth started in mid-April and continued until the end of November, with the maximum growth in June and July. In trees with the removal of cork, leaf abscission occurred a little earlier and new branches had on average 23% fewer new leaves. The radial growth of the trees and its general seasonal pattern were not affected by cork removal in the year of removal or in the year after. The only observation was a shift of the maximum radial growth rate from June to July for the trees where cork had been removed.     

Occurrence of Phytophthora cinnamomi in cork oak forests in Italy

An increasing decline and mortality of cork oak trees have been recently observed in central Italy and Sardinia Island. Following surveys conducted in three declining cork oak forests, a Phytophthora species was consistently isolated from soil samples collected from trees displaying different level of decline. Based on morphological features, growth rates at different temperatures and analysis of DNA sequences of the ITS region, all isolates were identified as Phytophthora cinnamomi Rands. This pathogen caused large brownish lesions on inoculated freshly cut branches of cork oak. It was re‐isolated from all infected tissues. These findings represent the first report of P. cinnamomi on cork oak trees in Italy.     

Cellular structure and chemical composition of cork from the Chinese cork oak (Quercus variabilis)

Quercus variabilis Blume, the Chinese cork oak, is an oak species with a thick cork outer bark. The cork is exploited at a limited scale in China and considered of lower quality than the commercial cork from Quercus suber. We studied an industrial cork granulate feedstock of Q. variabilis in relation to cellular structure and chemical composition and compared it to Q. suber cork under a material’s perspective. The cork of Q.variabilis has 1.1 % ash, 9.6 % extractives, 34.8 % suberin and 19.1 % lignin. The monosaccharide composition with shows a predominance of hemicelluloses: glucose 42.8 % of total neutral sugars, xylose 27.5 %, arabinose 15.4 %, galactose 9.0 %, mannose 4.0 %, rhamnose 1.2 %. The FT-IR spectrum shows the indicative peaks of suberin. The composition is overall similar to that of Q. suber cork. Q. variabilis cork has the typical cellular characteristics of bark cork tissues with a regular and radially aligned structure of cells without intercellular voids. Solid volume fraction was estimated at approximately 16 %. Compared with Q. suber, the Q. variabilis cork cells are smaller, the cell wall undulation and the overall row alignment less homogeneous, the cell surface is irregular and the solid volume proportion higher. The characteristics of Q. variabilis cork support its use as a cellular material for sealing, insulation and energy absorption, but the overall quality is lower than that of Q. suber cork. The negative impact of the higher density and structural lower uniformity at tissue and cell level should be evaluated for processing and product performance.     

Optimizing the debarking and cutting schedule of cork oak stands

Key message

Optimal management of cork oak forest stands was analyzed for different site indices and cork growth rates. Optimal debarking intervals varied during the rotation and were sometimes shorter or longer than the officially recommended range of 9–14 years.

Context

Quercus suber L. is one of the most important multipurpose tree species in the Mediterranean area. Its main product is cork, appreciated for its elasticity, impermeability, and thermal insulation properties. Cork oaks are debarked at constant intervals, which vary from 9 to 14 years depending on the area. However, since the growth rate of cork is not constant during the rotation, it may be optimal to use variable debarking intervals.

Aims

This study optimized the debarking and cutting schedules of Quercus suber stands and analyzed the influence of economic and stand-related factors on optimal management.

Methods

The study employed a simulation system where the existing growth and yield models for Quercus suber were used with a non-linear derivative-free optimization algorithm. Discount rates and cork prices were tested as economic factors and cork growth rate and site productivity as stand-related factors.

Results

The optimal debarking interval varied during the rotation. Increasing cork growth rate increased the optimal number of debarkings and shortened their interval. Decreasing discount rate increased the optimal number of debarkings during rotation while decreasing cork price decreased the number of debarkings.

Conclusion

The profitability of the management of cork oak stands depends on site fertility and stand density; management is not profitable on poor sites or at high discount rates. This study is the first that simultaneously optimizes the cutting and debarking schedule of cork oak stands, allowing the debarking interval to vary.

     

Differences of raw cork quality in productive cork oak woods in Sicily in relation to stand density

Differences of raw cork quality, in terms of thickness and porosity, were observed in productive cork oak (Quercus suber L.) stands in the Nebrodi and Iblei mountains of Sicily in relation to stand density. Five study areas were chosen across a range of different stand density satisfying specific sampling requirements among two managed cork oak woods.Analysis of variance using Fisher–Snedecor‘s F test (p < 0.05) were used to identify statistically significant differences of cork quality and sylvicultural, dendrometric and cork stripping parameters between study areas within each stand. Relationships between cork quality and sylvicultural and dendrometric parameters were also tested using Pearson's correlation coefficient (r). Results shown that sylvicultural practices are a key factor influencing cork quality: statistically significant differences in cork thickness and crown area were observed between areas at different stand density; positive correlations were found between cork thickness and crown area in lower density study areas favouring a significant vegetative activity; negative correlations were found between parameters of cork stripping (cork stripping coefficient, cork stripping surface, maximum height of cork stripping) and cork thickness. No correlations were found between cork porosity and dendrometric or sylvicultural parameters, demonstrating the probable genetic origin of cork porosity. Results shown that cork quality parameters and stand attributes may be used in sustainable management models of cork oak forests.     

Birds as predators of cork and holm oak pests

The recent decline of Mediterranean oak woodlands in SW Iberian Peninsula is related to insect pests which affect both cork oak (Quercus suber) and holm oak (Quercus rotundifolia). We identified twenty-six bird species as potential regular predators of twenty major pests by reviewing the diet of breeding, wintering and resident species in this ecosystem. Foraging guilds are strongly associated with predation at distinct stages of the pests’ life-cycle: ground-foragers prey on overwintering pupae and larvae of seed-borers, tree-foragers prey on eggs, larvae and pupae of defoliating and wood-boring pests, and aerial-sweepers prey on airborne imagines. Bird predation can cover the complete life-cycle of pest species because different species may be complementary due to a dissimilar exploitation of foraging niches and periods. Small generalist tree-foraging passerines are important pest predators given their high densities and widespread distribution in Mediterranean oak woodlands, but management practices can have a significant negative effect in their populations.     

Pathways for resilience in Mediterranean cork oak land use systems

Context

Loss of woodlands and degradation of vegetation and soil have been described for all Mediterranean-type ecosystems worldwide. In the Western Iberian Peninsula, overexploitation of evergreen cork oak land use systems has led to soil erosion, failures in oak recruitment, and loss of forests. Degraded and dry sites are quickly colonised by pioneer heathland rockrose (Cistus spp.) shrubs forming highly persistent patches.

Aims

Although traditionally shrublands have been considered as a transient successional state, we present evidence that they can represent persistent alternative states to former cork oak forests.

Review trends and conclusions

We first describe how Mediterranean vegetation evolved in the Iberian Peninsula and the role of fire and long-term human management as main disturbances. We then discuss alternative pathways through state-and-transition models indicating the ecological and land use variables that halt cork oak regeneration and recruitment and drive vegetation transitions towards persistent shrublands. Unless concerted management actions and restoration programmes are undertaken, the cork oak land use systems will not be sustainable.     

Establishment of cryopreserved gene banks of European chestnut and cork oak

Cryopreservation of selected genotypes of European chestnut and cork oak was carried out in two laboratories in a project involving conservation of field collections. Plant material was selected on the basis of disease resistance (chestnut), growth habit, phytosanitary performance and cork quality (cork oak). The cryopreservation technique comprised of vitrification of shoot apices isolated from in vitro stock shoot cultures (chestnut) and somatic embryos (cork oak). Forty-three out of 46 chestnut genotypes assayed survived the freezing process, but only 63% recovered their capacity to produce new shoots. After completion of multiplication and rooting steps, the surviving shoots produced plants that were morphologically identical to those derived from non-supercooled material. All 51 cork oak genotypes withstood freezing and were able to produce new somatic embryos through a process of secondary embryogenesis. Multiplication and germination of the recovered embryos enabled production of plants that were morphologically identical to those derived from non-supercooled material. In light of the results obtained, long-term cryopreservation of these species is feasible, thereby ensuring conservation of valuable genotypes during field evaluation.     

Radial variation of wood density components and ring width in cork oak trees

The radial variation of ring width and wood density was studied in cork oaks (Quercus suber) using microdensitometry. The observations were made in young never debarked cork oaks (30–40 years of age) and in mature trees under cork production (37–60 years of age). The cork oak wood is very dense (mean ring density 0.86 g.cm^?3, between 0.79 g.cm^?3 and 0.97 g.cm^?3) with a small intra-ring variability (mean earlywood density 0.80 g.cm^?3 and latewood density 0.90 g.cm^?3). The density components decreased from pith to bark more rapidly until the 15th ring, and then only slightly. There were no significant differences in the mean density components between never debarked trees and trees under cork production but their outwards decrease was accentuated in the never debarked trees. The annual growth was high, with a ring width mean of 3.9 mm (4.2 mm in the first 30 years) and the latewood represented 57% of the annual growth.     

Long-term understory recovery after mechanical fuel reduction in Mediterranean cork oak forests

In the Euro-Mediterranean region, mechanical fuel reduction is increasingly used in response to the mounting occurrence of catastrophic wildfires, yet their long-term ecological effects are poorly understood. Although Mediterranean vegetation is resilient to a range of disturbances, it is possible that widespread fuel management at short intervals may threaten forest structural complexity and the persistence of some plant species and functional types, with overall negative consequences for biodiversity. We used a chronosequence approach to infer woody vegetation changes in the first 70 years after understory clearing in upland cork oak (Quercus suber) forests, and to assess how these are affected by treatment frequency. Across the chronosequence there was a shift between plant communities with contrasting composition, structure and functional organization. Understory cover increased quickly after disturbance and a community dominated by pioneer seeder and dry-fruited shrubs (Cistus ladanifer, C. populifolius, Genista triacanthos, and Lavandula stoechas) developed during about 15 years, but this was slowly replaced by a community dominated by resprouters and fleshy-fruited species (Arbutus unedo, Erica arborea) >40 years after disturbance. During the first 15 years there were rapid increases in woody species richness, vertical structural diversity, cover by Q. suber juveniles and saplings, and shrub cover at <1.5 m strata, which levelled off or slightly declined thereafter. In contrast, tree species richness, tree density and density of arboreal A. unedo and E. arborea, vertical structural evenness, and cover at >1.5 m strata increased slowly for >50 years. Treatment frequency showed strongly negative relationships with species richness, structural diversity and evenness, and horizontal and vertical understory cover, particularly that of slowly recovering species. These findings suggest that fuel reduction programs involving widespread and recurrent understory clearing may lead to the elimination at the landscape scale of stands with complex multi-layered understory occupied by large resprouters and fleshy-fruited species, which take a long time to recover after disturbance. Fuel management programs thus need to balance the dual goals of fire hazard reduction and biodiversity conservation, recognizing the value of stands untreated for >50 years to retain ecological heterogeneity in Mediterranean forest landscapes.     

Thermal optima of photosynthetic functions and thermostability of photochemistry in cork oak seedlings

Temperature effects on photosynthesis were studied in seedlings of evergreen Mediterranean cork oak (Quercus suber L.). Responses to changes in temperature and the temperature optima of maximal carboxylation rate (V(cmax)) and maximal light-driven electron flux (J(max)) were estimated from gas exchange measurements and a leaf-level photosynthesis model. The estimated temperature optima were approximately 34 and 33 degrees C for V(cmax) and J(max), respectively, which fall within the lower range of temperature optima previously observed in deciduous tree species. The thermostability of the photosynthetic apparatus was estimated according to the temperature at which basal chlorophyll a fluorescence begins to increase (T(c)). The T(c) was highly variable, increasing from 42 to 51 degrees C when ambient temperature rose from 10 to 40 degrees C, and increasing from 44 to 54 degrees C with decreasing soil water availability while net CO(2) assimilation rate dropped to almost zero. When a heat shock was imposed, an additional small increase in T(c) was observed in drought-stressed and control seedlings. Maximal T(c) values following heat shock were about 56 degrees C, which, to our knowledge, are the highest values that have been observed in tree species. In conclusion, the intrinsic temperature responses of cork oak did not differ from those of other species (similar T(c) under ambient temperature and water availability, and relatively low thermal optima for photosynthetic capacity in seedlings grown at cool temperatures). However, the large ability of cork oak to acclimate to drought and elevated temperature may be an important factor in the tolerance of this evergreen Mediterranean species to summer drought and high temperatures.     

Opportunity for hybridization between two oak species in mixed stands as monitored by the timing and intensity of pollen production

The opportunity of cross-pollination in mixed stands of two oak species (cork oak and holm oak) was studied by characterizing individual phenologies of flowering. In the spring of 1998 at one stand consisting of 64 marked trees, there was a period of 19 days when maximal pollen release in one species and stigma receptivity in the other occurred simultaneously, enabling interspecific gene flow in either direction. This happened in spite of an average time separation of 22 days between the two species, reflecting a considerable intraspecific variation in the timings of flowering. Flowering intensities (as estimated from male flower abundance) were high, but fruiting intensities were comparatively low. Shortly after pollination, considerable abortion of female flowers and early fruits was recorded. In 2000, the interspecific overlap of phenologies was drastically reduced due to a delay in cork oak flowering. On the other hand, the individual timings were repeatable for most trees, at least in holm oak. Two other mixed stands were subject of parallel studies, with similar results in all traits except for a less dramatic reduction in fruiting intensities. In spite of the high opportunity for cross-pollination in 1998, and given the lack of hybrids among the progenies from the subsequent fruiting season [Oliveira, P., Custódio, A.C., Branco, C., Reforço, I., Rodrigues, F., Varela, M.C., Meierrose, C., 2003. Hybrids between cork oak and holm oak: isoenzyme analysis. Forest Genet. 10, 283–298], it can be concluded that the prerequisite of cross-pollination is clearly insufficient for hybridization to succeed. Post-pollination processes must play an important role in the maintenance of reproductive isolation between the two species.     

Influence of site factors on the impact of Phytophthora cinnamomi in cork oak stands in Portugal

Although decline of cork (Quercus suber) and holm oak trees (Quercus rotundifolia) has been described in Portugal in the late years of the 19th century, its development has become a motive of high concern during the last two decades. The presence of Phytophthora cinnamomi in cork and holm oak stands was surveyed in four different regions of the country (Trás‐os‐Montes, Alentejo, Ribatejo and Algarve) during 1995–98. Tree decline severity, sudden death and site characteristics were assessed in 56 sites representing varied conditions. The pathogen was isolated from oak roots and rhizosphere samples in 27 of those places. Other plant species from natural vegetation were sampled in three active disease centres. This survey showed that 56% of the surveyed species of shrub flora were infected with P. cinnamomi, which was detected mainly on species belonging to the families Ericaceae, Cistaceae and Leguminosae. Recovery of P. cinnamomi was more frequent in shallow soils (Leptosols and complex Leptosols and Luvisols). These soils are more common in the south (Algarve), where decline has a high impact. Soils with low fertility and low mineral nutrient levels, particularly phosphorus, seemed to favour infection. Site aspect and topographic tree situation were also evaluated. Sites facing south showed higher occurrence of P. cinnamomi, which was also more frequent in slopes and valleys than on hilltops. In Algarve, a relationship could be established between the crown status and the presence of P. cinnamomi in roots and rhizosphere. Different morphotypes of P. cinnamomi could be distinguished in vitro, and their occurrence in the field was correlated with particular site characteristics. Further research needs and management strategies to limit the extension of the disease are discussed.     

Vessel size and number are contributors to define wood density in cork oak

Cork oak (Quercus suber L.) has a dense wood that allows high-quality uses. In the present work, we study the influence of vessel characteristics, measured through image analysis and optical microscopy, on wood density, measured using X-ray microdensitometry, on 40-year-old trees. Vessel area increases with cambial age (5403–33064 μm²), while wood density decreases (1.229–0.836 g/cm³). The number of vessels is relatively constant at 6 vessels/mm², while vessel proportion in cross-section increases from 3.3% near the pith to 20.5% near the bark. In growth rings closest to the pith, with high wood density and low vessel area, the relationship between the two variables is linear (R ² = −32.1%, P < 0.01) but with increasing tree age and vessel size, the wood density remains rather constant, suggesting that decreases in density might compromise mechanical support of the tree at a stage when the increase in cross-sectional area alone might not provide mechanical stability. Other anatomical characteristics not considered in this study, like large xylem rays that increase with cambial age, may be responsible for the constant density.