Can we use shelterwoods in Mediterranean pine forests to promote oak seedling development?

The use of shelterwoods to favour the development of natural or underplanted seedlings is common in temperate forests but rare in the pine forests of the Mediterranean area. Our aim was to assess the use of shelterwoods in Aleppo pine (Pinus halepensis) woodlands in southern France to promote the survival and growth of two co-occurring oak species: the deciduous Quercus pubescens and the evergreen Quercus ilex.Twelve Aleppo pine stands were selected and differentially thinned to create a light shelterwood (basal area = 10 m²/ha, irradiance 52%), a medium shelterwood (basal area = 19 m²/ha, irradiance 33%) and a dense shelterwood (basal area = 32 m²/ha, irradiance 13%). A total of 1248 sowing points, half composed of Q. pubescens and half of Q. ilex, were then set up in these three conditions. Seedling survival and growth were monitored for 3 years. Plant stress was assessed by measuring predawn leaf potential and photosynthetic performance through the F_v/F_m ratio. Soil moisture was also recorded at two depths during two growing seasons.Survival was high for both species in all three conditions due to three consecutive wet years. The lowest survival was recorded for Q. pubescens in the dense shelterwoods. Growth in diameter and height increased from the dense to the light shelterwoods. Shrubs developed more strongly in the light shelterwood, and increasing shrub cover enhanced height growth. Photosynthetic performance was lowest for Q. pubescens in dense shelterwoods and highest in light shelterwoods, whereas the reverse was true for Q. ilex. The lowest predawn potentials were recorded in the dense shelterwoods even though higher soil water content values were measured in this treatment during the summer drought.We show that light shelterwoods were more beneficial to growth than denser ones, indicating control mainly by light availability during the 3 years of the study. However, as lower soil moisture at 30-50 cm depth and faster understorey development were also recorded in this condition, more extended observation is needed to determine whether this benefit persists in subsequent years.     

Trends in evergreen oak suitability from assembled species distribution models: assessing climate change in south-western Europe

The expected changes to the environmental conditions have concerned the scientific community over the last few decades. A rise in the mean temperature and a variation in rainfall patterns could modify the current distribution of plant species. In this study, we analysed four evergreen oaks (Quercus ilex subsp. ilex, Q. ilex subsp. ballota, Q. suber and Q. coccifera) by means of species distribution models. Three algorithms were used: maximum entropy, logistic regression and environmental distance. Taxa occurrences were taken, chiefly from the National Forest Inventories, and climate data was retrieved from the WorldClim 1.4 project. The present period and four future scenarios were studied. The latter were carried out by averaging thirteen global circulation models (GCMs). Area under the curve was used for validating the models. Maps indicating the suitability and cosuitability among the evergreen oaks were developed. The potential distribution of evergreen oaks in the present period was found to be wider than the actual distribution. Simulations indicate that climate change would increase the cosuitability of western temperate areas for Mediterranean oaks. The use of different algorithms and GCMs, as well as the high validation values obtained, make the study robust. Oaks are an important source of income, especially Q. ilex subsp. ballota and Q. suber. Our findings contribute to our understanding of the dynamics of oaks, and can be considered for management programmes aimed at conserving this natural heritage.     

Leaf-level responses to light in two co-occurring <Emphasis Type="Italic">Quercus</Emphasis> (<Emphasis Type="Italic">Quercus ilex</Emphasis> and <Emphasis Type="Italic">Quercus suber</Emphasis>): leaf structure,chemical composition and photosynthesis

We studied morphological, biochemical and physiological leaf acclimation to incident Photon-Photosynthetic-Flux-Density (PPFD) in Quercus ilex (holm oak) and Quercus suber (cork oak) at Mediterranean evergreen oak woodlands of southern Portugal. Specific leaf area (SLA) decreased exponentially with increasing PPFD in both species. Q. ilex had lower SLA values than Q. suber. Leaf nitrogen, cellulose and lignin concentration (leaf area-based) scaled positively with PPFD. Maximum rate of carboxylation (V_cmax), capacity for maximum photosynthetic electron transport (J_max), rate of triose-P utilization (V_TPU) and the rate of nonphotorespiratory light respiration (Rd) were also positively correlated with PPFD in both Quercus species, when expressed in leaf area but not on leaf mass basis. Q suber showed to have higher photosynthetic potential (V_cmax, J_max^m and V_TPU^m) and a higher nitrogen efficient nitrogen use than Q.ilex. Leaf chlorophyll concentration increased with decreasing PPFD, improving apparent quantum use efficiency (Φ) in both Quercus species. We concluded that, in Q.ilex and Q.suber, leaf structural plasticity is a stronger determinant for leaf acclimation to PPFD than biochemical and physiological plasticity.     

Pathogenicity of three Phytophthora spp. causing late seedling rot of Quercus ilex ssp. ballota

Within a research project on quality of plants for forestation of agricultural lands, we studied the aetiology of a late seedling rot affecting holm oak (Quercus ilex ssp. ballota) in two forest nurseries in southern Spain. Major disease symptoms were foliage wilting and necrosis of feeder roots. Phytophthora cinnamomi, Phytophthora cryptogea and Phytophthora drechsleri were isolated from necrotic roots of holm oaks. Selected isolates of the three Phytophthora species were pathogenic to Quercus ilex ssp. ballota and Quercus suber seedlings in artificial inoculations. Soil flooding conditions were essential for infection and root rot development. There was no host specificity among the species, the isolates of P. cinnamomi being the most virulent in all inoculated plants. In these inoculations, Q. ilex ssp. ballota plants were more susceptible than those of Q. suber. This work is the first report of P. cinnamomi, P. drechsleri and P. cryptogea affecting Q. ilex ssp. ballota in forest nurseries.     

Fingerprinting and genetic variability in cork oak (Quercus suber L.) elite trees using ISSR and SSR markers

Quercus suber L., is a socially, economically and ecologically important forest species in rural areas of the Mediterranean basin. Fifty three elite-trees from nine stands of four provenance regions in the Community of Extremadura (Spain) were analysed with the aim to establish their DNA-fingerprinting and the genetic relationships among them. Two types of molecular markers, microsatellites and intermicrosatellites, were used for tree genotyping. Microsatellite markers could distinguish 94% of the trees. Observed and expected heterozygosities, and effective number of alleles were correlated with the polymorphism information content (PIC). Intermicrosatellite patterns were used to construct a dendrogram. They showed high levels of genetic diversity in these elite trees, without any clear relationship with provenance regions. Both types of markers detected a high level of intra-stand variability, which is common in Quercus species, probably due to its mating system, the low distance among stands and the small size of stands. Others factors that could affect this result, such as introgression between Q. suber L. and Q. ilex L., are also commented. The high level of genetic variability detected in these elite trees can be useful for improvement programs. Also the usefulness of SSR and ISSR markers to establish the DNA-fingerprinting of each tree could be focused to select clonal lines of commercial interest.     

Composition and structure of natural mixed-oak stands in northern and central Portugal

Stand composition and structure of natural mixed-oak stands of common-oak (Quercus robur L.) and pyrenean-oak (Quercus pyrenaica Willd.) were studied. Diverse compositional and structural elements in early and late successional stand stages were analysed. The study was conducted in north and central Portugal where different natural mixed oak forests types are located. The following mixed-oak forest types involving common-oak and pyrenean-oak were studied: common-oak & other hardwoods; common-oak & cork-oak (Quercus suber L.); ash (Fraxinus angustifolia Vahl) & pyrenean-oak; and pyrenean-oak & madrone (Arbutus unedo L.). Measurements were made in early and late successional stand stages on the different mixed oak forest types. Different stand characteristics and indices were used to describe and compare stand structure and composition. The study showed changes in species diversity and stand structure. Most tree species in mature stands are present in early stages but with higher abundance. Shannon diversity index may change between 0.798 and 1.915. Significant differences on species diversity and abundance were found depending on the forest type and successional stage. Mature mixed-oak forests have high species diversity with an abundance of small to medium tree size species. Species distribution and diameter differentiation indices range from 0.30 to 0.70 and 0.52 to 0.82, respectively, revealing significant structural complexity. The average number of standing and downed dead trees was 265 and 83 trees ha⁻¹ for early and late stage, respectively, with 6.9 and 65.4 m³ ha⁻¹. Higher values of stand diversity index were 41 and 53 in more complex and developed forests. Later stand stages have complex structure, with a wider range of tree diameter distribution and higher degree of irregularity.     

Root system of Quercus suber L. seedlings in response to herbaceous competition and different watering and fertilisation regimes

Different management practices are commonly applied to increase pasture yield of Mediterranean open woodlands, but the consequences of increasing competition for soil resources with these practices on tree recruitment are still unknown. In a greenhouse study, belowground competition of Quercus suber L. seedlings growing together with natural (Oak_NP) or improved pasture (Oak_IP) was evaluated, and their root systems compared with those of seedlings growing in bare soil (Oak_BS). Two watering levels and two regimes of P₂O₅ fertilisation were also tested. Because of competition, the Oak_IP seedlings had their fine root mass density, coarse root length, and shoot mass reduced by up to 40, 36, and 39%, respectively, when compared to Oak_NP seedlings. Oak_NP and Oak_BS seedlings showed similar average root density parameters and shoot mass values, indicating that Q. suber seedlings growing with natural pasture is a low competitive system. High availability of water and phosphorus did not mitigate the strength of competition between herbaceous plants and oak seedlings, and favoured the pasture to the detriment of the trees. Our findings suggest that P₂O₅ fertilisation and irrigation practices performed to improve herbaceous productivity will negatively influence recruitment of Q. suber seedlings.     

The impact of atmospheric deposition and climate on forest growth in European monitoring plots: An individual tree growth model

In the climate change discussion, the possibility of carbon sequestration of forests plays an important role. Therefore, research on the effects of environmental changes on net primary productivity is interesting. In this study we investigated the influence of changing temperature, precipitation and deposition of sulphur and nitrogen compounds on forest growth. The database consisted of 654 plots of the European intensive monitoring program (Level II plots) with 5-year growth data for the period 1994–1999. Among these 654 plots only 382 plots in 18 European countries met the requirements necessary to be used in our analysis. Our analysis was done for common beech (Fagus sylvatica), oak (Quercus petraea and Q. robur), Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). We developed an individual tree growth model with measured basal area increment of each individual tree as responding growth factor and tree size (diameter at breast height), tree competition (basal area of larger trees and stand density index), site factors (soil C/N ratio, temperature), and environmental factors (temperature change compared to long-term average, nitrogen and sulphur deposition) as influencing parameters. Using a mixed model approach, all models for the tree species show a high goodness of fit with Pseudo-R² between 0.33 and 0.44. Diameter at breast height and basal area of larger trees were highly influential variables in all models. Increasing temperature shows a positive effect on growth for all species except Norway spruce. Nitrogen deposition shows a positive impact on growth for all four species. This influence was significant with p < 0.05 for all species except common beech. For beech the effect was nearly significant (p = 0.077). An increase of 1 kg N ha⁻¹ yr⁻¹ corresponds to an increase in basal area increment between 1.20% and 1.49% depending on species. Considering an average total carbon uptake for European forests near 1730 kg per hectare and year, this implies an estimated sequestration of approximately 21–26 kg carbon per kg nitrogen deposition.     

Effects of resource availability on condensed tannins and nitrogen in two Quercus species differing in leaf life span

Seedlings of Quercus ilex and Q. cerrioides, an evergreen and a winter-deciduous oak co-occurring in western-Mediterranean forests, were grown at two light regimes (8 and 36% of photosynthetically active radiation), at two water regimes (500 and 800 mm) and with two nutrient availabilities (standard substrate and 7% increase in soil N). The concentrations of soluble condensed tannins (CT) and nitrogen in the leaves were analyzed to test the phenotypic plasticity of these commonly related parameters in two con-generic species with contrasting leaf habit. Q. ilex contains seven times more CT and a few less N than Q. cerrioides. Light increased CT, whereas neither fertilization nor water had an effect on CT. N concentration was decreased by light, increased by fertilization and not affected by water treatment. Plant growth was increased by light but not affected by fertilization or water treatment. CT were negatively correlated with N concentration. CT of the evergreen species exhibited greater plasticity than the deciduous one as reflected by a steeper negative correlation among nitrogen and CT concentrations in Q. ilex. Given the antiherbivory activity of CT, this implies that in less shaded environments, e.g. canopy aperture by disturbances, leaf tissue quality for herbivores will be much more reduced in Q. ilex than in Q. cerrioides. Higher leaf CT in Q. ilex and its higher plasticity to light availability may explain the higher browsing by sheep in Q. cerrioides than in Q. ilex resprouts, as well as the low recruitment rates of seedlings of the former species, reported in other studies.     

Effects of grazing exclusion and environmental conditions on the soil seed bank of a Mediterranean grazed oak wood pasture

Large seed banks in the soils of Mediterranean wood pastures can allow the composition of the understorey vegetation to adapt to changing conditions such as under-grazing, grazing exclusion and climate change. This three year study investigated the effect of grazing exclusion on the transient and persistent seed banks of 23 areas of a Mediterranean wood pasture of Quercus suber L., Q. ilex L. and Q. pubescens Willd. A canonical correspondence analysis was used to determine the effect of topo-climatic (elevation, aspect, slope, rainfall, temperature, tree coverage), soil (pH, soil texture, and soil nitrogen, phosphorus, lime and organic carbon content) and biodiversity (Shannon index, species richness index, and Pastoral Value) variables on the soil seed bank under grazed and ungrazed conditions. The size of the persistent seed bank increased with rainfall, grazing, and the available phosphorus content of the soil. Specific site by site grazing regimes could increase the abundance of legumes in the soil seed bank and the species richness and diversity of the understorey vegetation. These results can help guide the conservation management of this silvopastoral area.     

Litter flammability in oak woodlands and shrublands of southeastern France

Characterizing the flammability of litter fuels is of major importance for assessing wildland fire ignition hazard. Here we compared the flammability of litter within a mosaic of Quercus suber (cork oak) woodlands and shrublands in a Mediterranean fire-prone area (Maures massif, southeastern France) to test whether the characteristics and the flammability of litter vary with the vegetation types. We tested experimentally the ignitability, the sustainability, the combustibility and the consumability of undisturbed (=non-reconstructed) litter samples with a point-source mode of ignition. Although the frequency of ignition was similar between all the vegetation types, we distinguished four groups having litter of specific composition and flammability: low and sparse shrublands dominated by Cistus species, medium shrublands with cork oak, high Erica shrublands with sparse cork oak woodlands, and mixed mature oak woodlands with Q. suber, Q. ilex and Q. pubescens. As these vegetation types corresponded to a specific range of past fire recurrence, we also tested the effect of the number of fires and the time since the last fire on litter flammability. Litters of plots recurrently burned had low ability to propagate flames and low flame sustainability. We discuss how the recent fire history can modify vegetation and litter flammability, and thus the fire ignition hazard.     

Fire-vegetation interplay in a mosaic structure of Quercus suber woodlands and Mediterranean maquis under recurrent fires

We examined the effects of fire recurrence on a mosaic structure of Quercus suber and Erica-Cistus shrubland communities of Southeastern France in order to improve the understanding of fire-vegetation interplay. Plant communities that were similar in 1959 (woodlands on shrublands called maquis on acidic soils) were compared along a gradient of fire recurrence, from 0 to 4 fires, with different time intervals between fires. The results showed that understory cover increased roughly with fire recurrence, whereas tree height, cover, density, stand basal area and litter depth decreased. Different fire recurrences along the past decades led to different vegetation types. High fire recurrence corresponded to maquis and sparse cork-oak woodlands while pure oak woodlands established in the absence of fire during the same period. In all, the diversity of tree diameter decreased with fire recurrence. High fire recurrence (3 or 4 fires in 50 years) led to a simple vertical structure of vegetation with a mono-layered shrub cover and few Quercus suber trees. In contrast, spatial connections between plants were maximal at longer time intervals, leading to a multi-layered vegetation. We finally discuss the potential implications of past fires on the behavior of future fires in the perspective of a sustainable management of these Mediterranean ecosystems.     

Fire-driven oak regeneration in French Mediterranean ecosystems

Regeneration by seeds for cork oak (Quercus suber) and companion oaks (holm oak Quercus ilex and downy oak Quercus pubescens) is likely to be poor in the fire-prone Maures massif (southern France) but the causes are poorly known. Our objective was to assess the effective recruitment for these three oak species and their temporal pattern of recruitment, in order to determine the main limitation factors and the regeneration window of each species. We studied oak recruits (height <3 m) in naturally regenerated populations according to a gradient of fire recurrence and in five main vegetation types including shrublands and mixed mature woodlands. Fire recurrence was the main explanatory factor of oak recruitment, either directly or through vegetation type and microsite characteristics. The results indicate nil to low recruitment for holm oak and downy oak in shrublands, especially those recurrently burned and dominated by Cistus species. Cork oak recruited better than the other oaks in medium and high shrublands dominated by Erica arborea. In contrast, recruitment was high for holm and downy oak in mixed oak stands and mixed pine-oak stands that have not burned for decades. Microsite conditions such as coverage by litter and shrubs influenced oak recruitment, whereas landscape configuration and stand basal area had no influence. Our results suggest that strategic shrub-clearing, oak planting and protection of mixed oak woodlands as seed sources would help maintaining oak populations in the woodland–shrubland mosaic.     

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.

     

Fire resistance of tree species explains historical gallery forest community composition

Fire, climatic variability, and grazing by large herbivores have historically limited woody vegetation in the tallgrass prairie region of North America to gallery forests in protected areas along rivers and streams. Fire, in particular, has been a strong selective pressure against woody vegetation. Consequently, we expect that dominant tree species in these forests have developed mechanisms for tolerating periodic surface fires. Susceptibility of trees to fire damage depends in part on key properties of bark which influence heat transfer to the vascular cambium, including thickness, density, and moisture content. An historical (1983) survey of Konza Prairie Biological Station in northeast Kansas, USA indicated that gallery forests were co-dominated by Quercus macrocarpa and Quercus muehlenbergii, while Celtis occidentalis occurred as an important sub-dominant species. Populus deltoides, Gleditsia triacanthos, and Juniperus virginiana were relatively uncommon. To test the hypothesis that historically dominant gallery forest tree species are more resistant to fire damage than uncommon species, fire was applied to the bark of 10 individuals of each of these six species under conditions mimicking surface fires (400 °C for 120 s). Maximum temperature at the vascular cambium, bark thickness, bark moisture content, and bark density were measured. Trees were considered fire-resistant if the vascular cambium temperature remained below the thermal cell death threshold, 60 °C, throughout the treatment. Using logistic regression, bark thickness was found to be a significant predictor of lethal cambium temperatures (P = 0.002), while neither bark density nor moisture content were significantly related to lethal cambium temperature (P = 0.279 and P = 0.131, respectively). Across species, a minimum bark thickness of approximately 8.6 mm was necessary to maintain the vascular cambium temperature below 60 °C. Trees that produce thick bark quickly in juvenile size classes (P. deltoides, Q. macrocarpa, and Q. muehlenbergii) experienced lower temperatures at the vascular cambium than those which do not develop thick bark with increasing diameter (C. occidentalis, G. triacanthos, and J. virginiana). Ranking these tree species by either the DBH or age needed to develop the minimum protective bark thickness largely agreed with ranking based on historical relative importance. As fire frequency and intensity decrease in remnant tallgrass prairie of North America as a result of habitat fragmentation, fire suppression, and changing land management, fire-sensitive species may increase in relative importance in gallery forests because of increased juvenile survival.     

Diversification of Pinus halepensis forests by sowing Quercus ilex and Quercus pubescens acorns: testing the effects of different vegetation and soil treatments

The goal of this study was to develop management strategies favouring establishment and survival of holm oak (Quercus ilex L.) and downy oak (Quercus pubescens Willd.)??two species co-occurring in Southern France??in mature Allepo pine forests (Pinus halepensis Mill subsp. halepensis). An experimental design was assessed in a partially cut mature stand of Aleppo pine in which five soil and vegetation treatments??chopping, chopping followed by scarification in one or two directions, prescribed burning, control??and two slash treatments (presence/absence) were applied. A total of 1,600 sowing points, each composed of 3 Q. ilex or Q. pubescens acorns, were installed in the different treatments in November of two consecutive years at 6 and 18 months, after the end of treatments. Survival was monitored 3 and 2 years after sowing, soil surface at the sowing points was characterized at different dates, and predawn leaf water potentials were measured during the dry season. High mortality occurred after the first summer, but survival after 1 year was 2.3?C5.2 higher in Q. ilex than in Q. pubescens, confirming that Q. ilex was better adapted to the drier parts of the Mediterranean area. Survival was significantly influenced by the treatments, but there was a variable response between the two sowing years under most of the treatments. Only intense fire proved the most beneficial treatment for seedling survival in both years. The micro-local soil cover conditions induced by the treatments played a major role in explaining oak survival. In particular, grass cover (mainly Brachypodium retusum) proved to be largely unfavourable to seedling survival and growth, and this detrimental effect was also confirmed by lower predawn leaf water potential values with increasing grass cover. Acorn introduction designed to diversify mature Aleppo pine forest after soil and vegetation treatments therefore has to be considered for treatments that most efficiently impair the pre-existing competing grass cover such as prescribed high-intensity fire treatment.     

Variation in the date of budburst in <Emphasis Type="Italic">Quercus robur</Emphasis> and <Emphasis Type="Italic">Q. petraea</Emphasis> across a range of provenances grown in Southern England

The widely observed advance in spring budburst across a range of temperate forest species due to climatic warming has received considerable attention. This change in forest phenology has important implications for the choice of species and provenances currently being planted, which need to be suited to both current and future climatic conditions. Using a provenance trial in the south of England, this study assessed variation in the timing of budburst across 23 different European provenances of oak (Quercus robur L. and Q. petraea (Matt.) Liebl.) over 8 years of observations. The order in which the different provenances reached budburst was related to provenance source latitude: the southern provenances were always earlier than those from more northerly latitudes. The statistical technique partial least squares regression was used to identify critical periods of both chilling and warming. A General Linear Model and three-dimensional temperature response surfaces were used to analyse the temporal trends in budburst. There was a negative correlation between the date of budburst and mean daily air temperature in both the chilling and warming periods for all provenances, which was statistically significant for a majority. Spring warming had a larger effect on budburst than winter chilling with a mean spring temperature-driven advance of 3.61 days/°C (standard error = 0.17 days/°C) and mean winter period temperature-driven advance of 0.99 days/°C (standard error = 0.17 days/°C). Surprisingly, there was no statistically significant interaction between mean air temperatures during the chilling and warming phases on the date of budburst.     

Impact of stand density on water status and leaf gas exchange in Quercus ilex

Tree thinning reduces tree-to-tree competition and likely contributes to the improvement of tree water status and productivity in water-limited systems. In this study, we examined the importance of competition for water among Quercus ilex trees in open woodlands by comparing the water consumption and physiological status of trees located along stand density gradients which ranged from 10% (low density; LD) to 100% (high density; HD) of canopy cover. The study was carried out at two sites which differed in mean annual rainfall (506 and 816 L m⁻²; D_site and W_site, respectively). Predawn and midday leaf water potential (ψ_d and ψ_m, respectively) and CO₂ assimilation rate (A) were measured every two weeks from mid May to mid September, in eight trees located along a stand density gradient at each site. Sap flow and soil moisture were measured only at D_site. Sap flow was continuously recorded by sap flowmeters (constant heating method) installed in 12 trees along two stand density gradients. Soil moisture (?) was measured every 20 cm for the first meter and then every 50 cm up to 250 cm. Measurements were conducted in 18 soil profiles, 6 located in HD and 12 in LD (six beneath and six out the canopy). At W_site, differences among stand densities for ψ and A were very small and emerged only at the end of the dry season. At D_site, ψ (both predawn and midday), A, ?, and sap flow density were significantly higher in LD trees than in HD ones. At D_site, some water remained unused in the soil at the end of the dry season beyond the canopy in the LD areas, and trees did not experienced such an acute water deficit (ψ_d > −1 MPa) as the HD trees did (ψ_d < −3 MPa). Summer tree transpiration at the stand level (E_stand) tended to saturate with the increase of canopy cover. E_stand increases by 32% when canopy cover goes from 50% to 100%. Results confirmed that the increase of tree-to-tree competition with stand density was much more significant at dry sites. In these sites, tree thinning is recommended as a way to maintain tree functioning.     

Early seedling establishment of two tropical montane cloud forest tree species: The role of native and exotic grasses

Tropical montane cloud forest has been undergoing a drastic reduction because of its widespread conversion to pastures. Once these forests have been cleared exotic grasses are deliberately introduced for forage production. Exotic grass species commonly form monodominant stands and produce more biomass than native grass species, resulting in the inhibition of secondary succession and tree regeneration. The purpose of this study was to assess the effect of native vs. exotic grass species on the early establishment of two native tree seedlings (Mexican alder, Alnus acuminata and Jalapa oak, Quercus xalapensis) on an abandoned farm in central Veracruz, Mexico. Seedling survival and growth were monitored (over 46 weeks) in relation to grass cover and height, and available photosynthetic active radiation (PAR). More seedlings survived in the presence of the native grass Panicum glutinosum than those growing with the exotic grass Cynodon plectostachyus (92% vs. 48%). The causes of seedling mortality varied between species; Q. xalapensis was affected by herbivory by voles but mainly in the exotic grass-dominated stands, whereas A. acuminata seedlings died due to competition with the exotic grass. A. acuminata seedlings increased more in height in the exotic grass-dominated stands (102 ± 7.8 cm) compared to native grass-dominated stands (51 ± 4.7 cm). Grass layer height, cover and available PAR were correlated (Pearson; p < 0.05). In the exotic grass dominated plots, grass layer height was correlated with the relative height growth rates of Q. xalapensis (Pearson; p < 0.05). These results indicate that the exotic grass may be affecting tree regeneration directly (grass competition) and indirectly (higher herbivory). Passive restoration may occur once P. glutinosum dominated pastures are abandoned. However, when C. plectostachyus dominates, introduction of early and mid successional tree seedlings protected against vole damage is needed.     

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.