Viability of holm and cork oak seedlings from acorns sown in soils naturally infected with Phytophthora cinnamomi

The emergence and survival of pregerminated holm oak (Quercus ilex) and cork oak (Quercus suber) acorns from two ecologically different dehesas (Mediterranean open woodlands) were studied in two soils from these stands naturally infected with Phytophthora cinnamomi, and in the same soils previously sterilized in the autoclave. Phytophthora cinnamomi was consistently isolated from the radicles of all unemerged and all emerged but dead seedlings from the unsterilized substrates. Seedlings of holm oak were more susceptible to P. cinnamomi than those of cork oak. Mortality of holm oak seedlings was significantly different depending only on soil treatment (sterilized or unsterilized), and it was 100% in unsterilized soils, independent of acorn provenance and soil origin. Mortality of cork oak seedlings was significantly different depending on the acorn origin and soil treatment, and on the interactions acorn origin × soil origin and soil origin × soil treatment. The demonstrated high susceptibility of holm and cork oak young seedlings to P. cinnamomi could be a limiting factor in Mediterranean open woodlands (dehesas) not only in natural regeneration processes but also when reforestation by direct sowing is implemented.     

Differential effects of vegetation restoration in Mediterranean abandoned cropland by secondary succession and pine plantations on bird assemblages

Two contrasting trajectories for vegetation restoration in agricultural landscapes are secondary succession following cropland abandonment that can regenerate woodlands (passive restoration) and conversion of cropland to tree plantations (active restoration), which have mostly focused on pine species in the Mediterranean Basin. We compared the effects of these two contrasting trajectories of vegetation restoration on bird assemblages in central Spain. Vegetation structure differed in the two restoration trajectories, pine plantations attaining higher tree cover and height (31% and 4.1 m, respectively) but lower strata complexity than secondary shrubland and holm oak woodland (which attained 10% and 1.4 m of tree cover and height, respectively). Bird species richness differed in stands under active or passive restoration trajectories, the former collecting a higher total number of species (4.2 species per 0.78 ha plot) than the latter (3.5 species per plot). The number of forest species increased with vegetation maturity in both restoration trajectories, but especially in stands under active restoration. The occurrence of woodland generalist species increased and of species inhabiting open habitats decreased in actively restored stands, being some of these latter species of high conservation priority in the European context but relatively common at the regional level. Bird species inhabiting pine plantations had broader habitat breadth at the regional level than those inhabiting secondary shrublands and woodlands. Maximum regional density did not differ between both restoration trajectories, but it increased with development of the herbaceous layer only at the secondary succession trajectory. The relative importance of species of European biogeographic origin was higher in mature pine plantations (58.9% of total bird abundance) than in mature holm oak woodlands (34.4%), whereas that of Mediterranean species was considerably higher in the latter (40.1%) than in the former (20%). Bird assemblages of relatively small patches of pine plantations are unable to reflect the regional avifauna, in contrast with the relationships between local and regional assemblage characteristics that can be found in isolated natural forests. We conclude that programs of vegetation restoration should base upon a range of approaches that include passive restoration, active restoration with a variety of tree and shrub species, and mixed models to conciliate agricultural production, vegetation restoration and conservation of target species.     

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.     

Restoring silvopastures with oak saplings: effects of mulch and diameter class on survival,growth, and annual leaf-nutrient patterns

In Southwestern Spain, multifunctional silvopastoral systems consisting of pastureland and open oak woodlands are known as Dehesas. These, and other similar systems of the Mediterranean basin, are currently threatened by increasing intensive land use. As a consequence, oak regeneration is declining and is in need of adequate management and active restoration. Traditional restoration practices outplant one-year-old, nursery-produced oak seedlings grown in 250–350 cm³ containers, but establishment and growth results are typically poor. This work examines holm oak (Quercus ilex L. subsp. ballota (Desf.) Samp.) grown in a non-conventional container size (24 l) and age (6–7 years) with three mulch treatments (control, stone, and straw). In an open Dehesa of SE Spain, 106 oak saplings were planted in March 2010, and survival, diameter at breast height (DBH), and leaf-nutrient concentrations were analyzed. Forty months after planting, all treatments showed high survival (81 %) but only straw-mulched saplings differed significantly (94 %) from control (74 %). DBH increased over time but showed no significant differences among mulch treatments. Saplings with high initial DBH showed the greatest growth and change in DBH at the end of the study period. Leaf-nutrient concentrations changed significantly in the year following outplanting. Bi-monthly foliar nutrient concentration data show decreases in P, K, Zn, and B and sharp increases in Ca and Fe. In this work, we provide some evidence concerning the viability of non-conventional oak size for restoring, regenerating, or building up new agroforestry or silvopastoral systems. A combination of saplings with more than 10 mm of DBH and straw mulch is recommended.     

Making Tropical Succession and Landscape Reforestation Successful

Forests are dynamic ecosystems, spatial mosaics of localized disturbances and regenerating vegetation. Following forest conversion to human land uses, expanding forest cover follows four different modes of reforestation: (a) spontaneous natural regeneration; (b) assisted natural regeneration; (c) agroforestry; and (d) commercial tree plantations. Natural regeneration fundamentally depends on the alignment of ecological and social factors at landscape scales. A deeper socioecological understanding of reforestation in the tropics will lead to the development of effective policies and governance structures that enhance forest cover and ecosystem services, protect biodiversity, and improve forest-dependent livelihoods.     

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.     

Restocking with broadleaved species during the conversion of Tsuga heterophylla plantations to native woodland using natural regeneration

The removal of conifers planted during the twentieth century on sites that had been woodland for many centuries, with the intention of restoring native broadleaved species, is an important aim of forestry policy in Great Britain. Current guidance generally advocates gradual removal of plantation trees using continuous cover silviculture and restocking by natural regeneration, but methods are largely untested. This study investigated natural regeneration of trees and shrubs at sites where western hemlock had been established either pure or in mixture with native broadleaved species. Western hemlock had been cleared from 70% of the sites. There were c. 10?C300 stems ha^?1 providing canopy cover of 10?C90%; the predominant broadleaved canopy trees were ash, beech, birch and oak. The ground flora of all sites was species poor; bramble was the predominant vegetation type overall, but grasses, rushes and shrubs were relatively more abundant on open sites. A total of 15 native tree species were regenerating, birch was most common and at most sites there were few valuable broadleaved timber species. Although combined numbers of seedlings and saplings varied from 200 to 20,000 ha^?1, large areas of most sites had few regenerating trees and <10% of their area was becoming restocked with timber species. Relationships between seedling numbers and site characteristics were complex and varied with species, but there was a consistent positive relationship between the presence of nearby parent trees and the frequency and abundance of seedlings. The results indicate that the current reliance on natural regeneration may be unwise especially for those broadleaved species such as oak and beech which are valued for their timber.     

Grazing affects shoot growth,nutrient and water status of Quercus ilex L. in Mediterranean open woodlands

Context

Mediterranean open woodlands (dehesa) have faced a dual process of intensification and abandonment of grazing which has resulted in alteration of the understory vegetation.

Aims

We analysed the effects of land use changes on the physiological status of holm oak in different open woodlands (dehesa) in southern Iberian Peninsula.

Methods

In an area of extensive grazing, we selected six paired plots (one grazed, one abandoned) and grouped them by habitat types according to understory composition (nearly all monospecific Cistus ladanifer L. shrub or mixed shrub). Six plots of moderate and heavy grazing intensity were chosen within a settled area of livestock use. Shoot growth, macronutrient concentrations and water content were assessed in samples of holm oak leaves.

Results

Abandonment of grazing affected some nutrient concentrations and water content of holm oak leaves, but the effects were different according to habitat type. C. ladanifer shrub reduced N and P concentrations and water content while mixed shrub increased P concentration and water content. High grazing intensity improved shoot growth and leaf N and Mg concentrations.

Conclusion

Extensive grazing could be a useful management tool to enhance growth, nutritional and water status of holm oak in a habitat with limited resources such as Mediterranean open woodlands (dehesa).     

Does canopy pruning affect foliage-gleaning birds in managed cork oak woodlands?

Semi-natural cork oak woodlands are a biodiversity rich agro-silvo-pastoral system covering large areas in the Mediterranean. Canopies of adult oaks are often pruned, but nothing is known about the consequences of this treatment on biodiversity. We evaluated the impact of pruning on birds that forage in cork oak canopies in an area of southern Portugal. We characterized the use of trees by foraging birds with focal observations, and analysed the effects of pruning on density with point counts on pruned and control areas. As pruning reduces the foraging substrate for foliage gleaners, we predicted that these species would have lower densities in pruned areas. Pruning did not substantially affect overall bird density or species richness. However, as predicted, the density of species that foraged mostly by gleaning in the canopy tended to be lower in pruned areas, especially in winter when differences were statistically significant. In this season the combined density of foliage gleaners in the pruned stations was only half of that in controls. Pruning is also common in other managed Mediterranean woodlands that are important for birds, such as holm oak woodlands and olive groves, and foliage gleaners are likely to be affected in those too. The cumulative effects of pruning on all these habitats need to be assessed, but our results already indicate that pruning has negative consequences that should be properly considered in management decisions.     

Genetic variation of pedunculate oak (Quercus robur L.) in the urban woodlands of Helsinki

We present a genetic profile of pedunculate oak (Quercus robur L.) in three woodlands of the Helsinki region. We genotyped all mature trees and samples of seedlings to evaluate patterns of genetic variation and gene flow, and evaluate the long-term stability and natural regeneration of these stands. Although five microsatellite loci showed no significant differences in heterozygosity between generations in most locations, significant (p?per se does not interfere with gene flow over the distances represented here. Based on our findings, a minimal-maintenance program reliant upon natural regeneration of oak in the public parks of Helsinki seems plausible.     

A spatial distribution model of cork oak (Quercus suber) in southwestern Spain: A suitable tool for reforestation

Cork oak (Quercus suber) is an evergreen tree characterized by a thick bark, which grows in Mediterranean schlerophyllous forests. It is most prevalent in the southwestern Iberian Peninsula. Despite the potential of the province of Huelva (southwestern Spain) to maintain mature forests of cork oak, the tree has been severely depleted and most forests have either disappeared or are seriously threatened. This paper presents a spatial distribution model of cork oak for the province of Huelva with a view to determining the optimal areas for reforestation. The model draws on all available digital cartographic information with respect to cork oak distribution: topographic data (altitude, slope and orientation) were obtained from a Digital Terrain Model (20 m scale); rainfall, temperature and PET models were based on data collected from a network of meteorological stations; litologic data derive from the litologic map of Huelva (1:100,000). The result of this work is a mesh of points at a resolution of 100 m, sufficient to meet the needs of any kind of reforestation or management programmes in the area studied. Each point of this mesh contains the corresponding values for bioclimatic, topographic and litologic variables in a georeferenced data matrix. The independent variables responsible for cork oak distribution (binary dependent variables) were then identified by means of binary logistic regression analysis. North-facing slopes, abundant annual rainfall and litology were the main explaining variables. The spatial distribution model was produced by applying the formula obtained to spatial analysis software. This model is proposed as a basis for future reforestation plans, especially in those areas most affected by forest fires.     

Research needs for restoring seasonal tropical forests in Thailand: accelerated natural regeneration

Accelerated natural regeneration (ANR) is a relatively cheap method of reforestation, which encourages natural establishment of indigenous trees and shrubs. It requires a low input of labour, but a high input of ecological information. In this paper, the knowledge required to predict and manipulate the natural regeneration of seasonal tropical forest is reviewed and areas in need of further research are identified. Regeneration will be influenced by five groups of potentially limiting factors; site resources (soil and microclimate); competition with weeds; site disturbance; occurrence of established woody plants or their propagules; seed dispersal by wild animals and birds. This paper considers each of these, their interactions with seasons, and each other. Collation of existing information on these topics, combined with the suggested further research, should facilitate the creation of tools that will enable practitioners to judge the regeneration potential of sites and to select the most appropriate ANR techniques.     

Ethephon: a tool to boost gum arabic production from Acacia senegal and to enhance gummosis processes

Gum arabic production from Acacia senegal is lower in sub-humid areas than arid areas. Water stress is thought to be the reason for higher yields in arid areas. The application of ethephon is thought to mimic the effect of water stress in other plants. The objective of this study was to determine if the application of ethephon would increase the gum yields of Acacia senegal under sub-humid conditions in Cameroon. Trees receiving 40 or 120 mg ethephon were compared to controls in field experiments at a semi-arid and a sub-humid location in Northern Cameroon, over two seasons. Two provenances from drier areas (Sudan) were compared to the local one. In the first season, gum yield of the local provenance treated with ethephon was increased by 400–600 % compared to the untreated trees. Gum yield at the semi-arid location was 77, 313 and 214 g/tree with 0, 40 and 120 mg ethephon/tree, respectively, while at the sub-humid location, it was 30, 186 and 114 g/tree with 0, 40 and 120 mg ethephon/tree. However, in the second season, the effect of ethephon was not significant in the semi-arid area, whereas it was evident in the sub-humid area (up to 478 g/tree). Moreover, ethephon did not affect gum yield of provenances from drier areas (Sudan). This showed that the water-stress hypothesis has to be refined. The development of ethephon-based tapping systems is promising, but requires further studies with a wider range of environmental conditions and A. senegal provenances.     

Plant and carabid beetle species diversity in relation to forest type and structural heterogeneity

The aim of this study was to evaluate the influence of forest structure (mainly resulting from human uses) and forest type (the identity of the dominant tree species) on biodiversity. We determined the diversity of two taxonomical groups: the understory vegetation and the edaphic carabid beetle fauna. We selected eight types of forest ecosystems (five replicates or stands per forest type): pine (Pinus sylvestris) plantations of three age classes (10, 40 and 80 years since reforestation), an old-growth relict natural pine forest, and four types of oak (Quercus pyrenaica) stands: mature forests with livestock grazing and firewood extraction, mature forests where uses have been abandoned, “dehesa” ecosystems and shrubby oak ecosystems. The results obtained by a global PCA analysis indicated that both tree size and dominant species influenced the ordination of the 40 forest stands. In general, carabids were more sensitive to changes in forest heterogeneity and responded more clearly to the analysed structural variables than the understory vegetation, although the species richness of both groups was significantly correlated and higher in case of oak forests. Pine forest ecosystems were characterised by the lowest species richness for both taxonomical groups, the lowest plant diversity and by the lowest coefficients of variation and, consequently, low structural heterogeneity. As a result, it was very difficult to discriminate the effects of the spatial heterogeneity and the dominant tree species on biodiversity.     

Influence of soil properties and burial depth on Persian oak (<Emphasis Type="Italic">Quercus brantii</Emphasis> Lindl.) establishment in different microhabitats resulting from traditional forest practices

In seasonally dry environments such as the Zagros woodlands (Iran), severe drought stress and lack of appropriate management practices can cause failure of oak afforestation or reforestation. We investigated the effect of soil properties and burial depth on Persian oak (Quercus brantii Lindl.) establishment in different microhabitats resulting from traditional forest practices. Four microhabitats that were based on forest structure were considered for oak acorn seeding: (1) inside old sprout clumps (ISPC); (2) under the canopy of tall trees (UCTT); (3) outside the canopy of tall trees and sprout clumps; and (4) near recent stumps or sprout clumps. Acorns were seeded at two depths (2 and 5 cm), and seedling survival and growth variables were recorded for 4 years, together with soil chemical and biological attributes. Stepwise discriminant analysis showed that a combination of total soil nitrogen, cation exchange capacity, available phosphorus and potassium, litter depth, microbial quotient, metabolic coefficient, substrate-induced respiration and earthworm abundance was the best variables to characterise the microhabitats. With the exception of pH, bulk density and soil texture, these variables were higher in UCTT and ISPC than in the other microhabitats. Seedling emergence and survival were greater at a seed depth of 5 cm than at 2 cm. Seedling height and shoot, root and leaf biomasses were higher in the UCTT microhabitat compared to the other microhabitats and correlated positively with soil nutrients contents and most of the soil biological variables but negatively with soil bulk density. This study emphasised the role of microhabitats in creating a “canopy effect” producing favourable physical, chemical and biological soil conditions. In particular, large oak trees form islands of fertility and therefore are of key importance for successful seedling establishment in forests subjected to intense human activities.     

Impact of targeted sheep grazing on herbage and holm oak saplings in a silvopastoral wildfire prevention system in south-eastern Spain

Several wildfire prevention programs in southern Europe are currently using livestock grazing for the maintenance of fuelbreaks. This silvopastoral management is valued for being sustainable and effective in reducing fuel loads, but few studies have analyzed other impacts linked to fuelbreak grazing. This paper reports on an experiment performed within the wildfire prevention program in Andalusia (southern Spain) with the aim of clarifying and quantifying the effect of fuelbreak grazing on herbage biomass, ground cover, herbage species composition, and growth of holm oak saplings. The study site, located in a semiarid Mediterranean environment, was grazed by a shepherded sheep flock from February to June in three consecutive years at a similar stocking rate. Livestock consumed between 33 and 68 % of herbage production in the different years, and the greatest fuel reduction (remaining dry matter of 200?kg?ha^?1) was registered in Year 2, when rainfall and herbage production was lowest. Ground cover was significantly affected by grazing: on average, the percentage of bare soil increased three-fold, while herbage cover was reduced by a quarter. The botanical composition of herbage varied remarkably between years, but very little between Grazed and Non-Grazed areas within each year. Non-browsed holm oak saplings became progressively larger than browsed ones, differences only reaching clear statistical significance at the end of the three experimental years. At this time, the volume of browsed saplings was 47–56 % smaller than that of non-browsed holm oaks, even though the former had also grown significantly in the course of the experiment.     

Development of oaks (Quercus petraea (Matt.) Liebl.) emerged from bird-dispersed seeds under old-growth pine (Pinus silvestris L.) stands

In East Germany, there are a lot of areas covered by old pine stands. They are growing on soils, on which under natural conditions without anthropogenic impact mixed stands of oak (Quercus petraea (Matt.) Liebl.) and pine (Pinus silvestris L.), would form a natural forest. An important objective of ecological silviculture in these areas is to convert the pure pine stands into mixed oak–pine stands by using natural regeneration methods. A highly appreciated assistant in this connection is the European jay (Garrulus glandarius L.). The remarkable results of its work have been found by analyzing the natural regeneration in a few old pine stands in the forest district of Weißwasser in Saxony. Although mother trees of oak are very scarce there, natural regeneration of oak was found everywhere in the research area. The oak regeneration, undoubtedly created by the jay, amounted to at least 2000 oaks/ha. They were mainly distributed at random, while pine regeneration was aggregated in places where gaps in the pine canopy occurred. Oak regeneration was established much earlier than pine regeneration. Thus, oaks exceeded pines in age, height and diameter. The oaks were also superior to pines concerning height increment for the last three years. Thus, there is a good chance for oak to defend its prevailing role in the regeneration in the future. Probably, the next forest generation will be composed of oak and pine trees. The analysis of the quality of the oak regeneration shows that there is no substantial difference to artificially sown oak stands. This indicates that the European jay creates oak stands sufficiently both in number and quality.     

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.     

Structure of pollarded oak forest in relation to aspect in Northern Zagros,Iran

We analyzed the structure of pollarded oak forest and biometric indices of pollarded oak species in relation to aspect in northern Zagros forests, western Iran. A number of 319 circular plots (0.1 ha) were established using a systematic random method over the study area. In each plot, for all trees (diameter at breast height ≥5 cm) diameter was measured and tree species was recorded. Total height, trunk height, and major and minor diameter axis of the crown of two trees in each plot (nearest tree to the center of the plot and the largest tree in term of diameter) were measured. As the dispersion of slope and altitude classes in the study area were identical, the effect of these factors was assumed to be constant. To evaluate the effect of aspect on biometric indices of oak trees a comparison was used for each oak species separately. The results indicated that the forest species composition differed statistically significant in main aspects except for easterly and westerly aspects (P < 0.01). The diameter of similar oak trees was significantly different except for Lebanon oak in northerly and southerly and Gall oak in easterly and westerly aspects (P < 0.01), but there was no significant difference between the total height and crown area of similar oak species in different aspects. Differences in diameter, height, and crown area distributions showed a significant difference in main aspects. The basal area and tree density in northerly and southerly aspects were significantly different (P < 0.05).     

Landscape dynamics in endangered cork oak woodlands in Southwestern Portugal (1958–2005)

Cork oak landscape dynamics were assessed over a 47-year period (1958–2005) in an endangered region of southwest Portugal using a GIS approach. The area of cork oak woodlands was maintained during this period, but shifts due to land abandonment were evident leading to transformation of cork oak agriculture areas into woodlands and of cork oak woodlands into shrublands (average yearly change rates of 0.6 and 0.1%, respectively). The multi-temporal landscape analyses showed that expansion and regression rates of cork oak forests were similar (15 and 16 ha year⁻¹, respectively). The main factor determining oak woodlands expansion and regression was related to land-use changes, but slope, aspect and soil type were also significant factors. The substitution of agriculture lands and oak woodlands by shrublands has a determining role in periods of disturbance and recovery of these Mediterranean ecosystems.