Differences in fine-scale genetic structure and dispersal in Quercus ilex L. and Q. suber L.: consequences for regeneration of mediterranean open woods

Cork oak (Quercus suber L.) and holm oak (Q. ilex L.) are among the most important tree species (economically and ecologically) in the Western Mediterranean region, where they define unique open woods (created and maintained by man) known as 'dehesas' in Spain. However, these formations are under increasing threat due to the lack of regeneration. We have analysed spatial genetic structure in a mixed parkland; inferences about gene dispersal have also been performed, according to the isolation by distance model. Noticeable differences have been detected between the species, despite their similar ecological roles. Restricted effective dispersal leads to kin structures in cork oak, up to 70 m, while no genetic structure is observed in holm oak. Our results suggest a very effective dispersal for the latter, with a local historical gene flow estimated between 55 and 95 m. This is the first time regeneration of Mediterranean oak parklands has been assessed from a genetic perspective. Effective gene flow detected for holm oaks allows us to discount the risk of inbreeding over successive generations. Thus, regeneration of Q. ilex dehesas will just require action directed to help the settlement of the saplings (such as limiting grazing). However, in those cases where densities are too low, more intense forestation (such as plantation and/or establishment of appropriate shelter) will be needed. The 'density threshold' for initiating regeneration will probably be higher for cork oak, due to its more limited dispersal and minor full-light tolerance.     

Assessing the resilience of Mediterranean holm oaks to disturbances using selective thinning

Climate change will increase the frequency and the intensity of droughts in the Mediterranean region, likely reducing growth and increasing mortality of holm oaks (Quercus ilex), one of the most abundant species of Mediterranean forests. In water-limited systems such as those of the Mediterranean, carbon allocation patterns strongly favour belowground accumulation, especially in large subterranean structures called lignotubers. The resilience of these forests depends largely on the replenishment rate of these carbon reserves after disturbances. An experimental thinning, with two intensities (removal of 40% and 80% of basal area), was performed in 1992 in a holm oak forest at the Prades Experimental Complex of Catchments (NE Spain). In 2002, a second thinning was carried out in subplots within the former experimental 0.5 ha plots. Samples from the lignotubers of holm oak trees were analyzed for starch, and both mobile and immobile chemical components, in order to assess the resilience of holm oaks to repeated disturbances. Our results show that after 10 years, starch stocks in the lignotubers have only recovered to half their former values. Removing 40% of the basal area instead of 80% is suggested to be the better managing option for this kind of forests.     

Natural hybridisation between kermes (Quercus coccifera L.) and holm oaks (Q. ilex L.) revealed by microsatellite markers

Hybridisation between species of the genus Quercus is a common phenomenon as a result of weak reproductive isolation mechanisms between phylogenetically close species that frequently co-occur in mixed stands. In this study, we use microsatellite markers to analyse introgression between kermes ( Quercus coccifera L.) and holm ( Q. ilex L.) oak, two closely related taxa that frequently dominate the landscape in extensive areas in the Mediterranean region. All tested microsatellites amplified and were polymorphic in both kermes and holm oaks. Bayesian admixture analyses showed a good correspondence between each species and one of the two inferred genetic clusters. Five sampled individuals were a priori tentatively identified as hybrids on the basis of intermediate morphological characteristics, and it was confirmed that they also presented mixed genotypes. However, we also detected different levels of genetic introgression among morphologically pure individuals, suggesting that successful backcrossing and/or reduced phenotypic expression of genetic variance in certain individuals may have resulted in strong convergence towards a single species phenotype.     

Patterns of seedling recruitment in West-Mediterranean Quercus ilex forest influenced by canopy development

Abstract. Quercus ilex (holm oak) coppice forests belong to the most representative communities in the western part of the Iberian Peninsula. Due to the high sprouting potential of holm oak after coppicing, the dynamics of recruitment of new individuals in existing populations has been largely overlooked. In these forests, the density of seedlings increases in old stands, as a result of the remarkable increase in the recruitment of 1-yr and 2-yr old seedlings, probably due to better environmental conditions for germination. Older seedlings show a tendency towards increasing survival under some degree of canopy closure, since their density appears to be higher just before complete canopy closure takes place. Nevertheless, the reduced growth of seedlings in older stands due to low light levels causes age distributions of holm oak seedlings to change drastically, with saplings older than 15 yr becoming very unusual in these stands. Therefore, a bottleneck of recruitment appears at this sapling stage and prevents vigorous regeneration just before thinning. Present conditions of genet density and management tend to prevent population renewal. Dynamics of the seedling bank reinforce the role of gap formation through small or large scale perturbations in population turnover.     

Trunk injection of fosetyl‐aluminium controls the root disease caused by Phytophthora cinnamomi on Quercus ilex woodlands

In Spain, Quercus open woodlands are animal ranching systems of organic production seriously threatened by the exotic pathogen Phytophthora cinnamomi. The root disease it causes kills thousands of oaks annually. Effective disease management needs to integrate different techniques, and the use of a resistance inducer such as fosetyl‐Al can play a key role, because the use of potassium phosphite is prohibited in Spain. In a woodland where the pathogen recently arrived, 60 holm oaks in three different defoliation classes (asymptomatic, slight and moderate defoliation) were selected for trunk injection with pressurised capsules containing 4% of commercial fosetyl‐Al or water (controls). Holm oaks were checked periodically for defoliation and presence of the pathogen in roots and rhizosphere soil. Three years after treatments, defoliation was significantly lower in oaks treated with fosetyl‐Al, which even increased canopy cover, in comparison with control oaks, independent of the initial defoliation class considered. Chlamydospore density in rhizosphere soil, as well as the presence of the pathogen into the roots, was not significantly influenced by fosetyl‐Al treatments, although a trend to a lower presence of P. cinnamomi in roots was observed in treated oaks at every soil inoculum density detected. This study has shown that fosetyl‐Al, a phosphonate registered as a fungicide in the European Union, provides protection to holm oaks against P. cinnamomi, even exhibiting a therapeutic effect on pre‐existing infections. Consequently, this effective measure should be considered as part of the integrated approach to control this highly destructive pathogen in holm oak woodlands.     

A global change-induced biome shift in the Montseny mountains (NE Spain)

Shifts in plant species and biome distribution in response to warming have been described in past climate changes. However, reported evidence of such shifts under current climate change is still scarce. By comparing current and 1945 vegetation distribution in the Montseny mountains (Catalonia, NE Spain), we report here a progressive replacement of cold‐temperate ecosystems by Mediterranean ecosystems. Beech (Fagus sylvatica) forest has shifted altitudinally upwards by ca. 70 m at the highest altitudes (1600–1700 m). Both the beech forests and the heather (Calluna vulgaris) heathlands are being replaced by holm oak (Quercus ilex) forest at medium altitudes (800–1400 m). This beech replacement has been observed to occur through a progressive isolation and degradation of beech stands. In ‘isolated’ (small and surrounded by holm oaks) beech stands, beech trees are 30% more defoliated, beech recruitment is 41% lower, and holm oak recruitment is three times higher than in ‘continental’ (large and continuous) beech stands. The progressively warmer conditions, complemented by the land use changes (mainly the cessation of traditional land management) are the apparent causes, providing a paradigmatic example of global change affecting distributions of plant species and biomes.     

Common cranesGrus grus and habitat management in holm oak dehesas of Spain

Changes in agricultural policies have favored tree clearing and removal of shrubs to favor intensive farming in the holm oakQuercus ilex dehesas of Iberia. The effect of these changes for bird species wintering in this agricultural landscape has been rarely analyzed. Here the effects are studied of farming changes in traditional holm oak dehesas on population size, social structure and time budget of common cranesGrus grus in eight traditional wintering sites in Spain. The role of acorn availability, the principal winter food of the species, in relation to management is also assessed. The number of cranes in each wintering area was not related to degree of agricultural intensification, nor to the availability of acorns. However, crane flock size decreased, and the relative occurrence of isolated family groups increased in less intensively transformed wintering areas characterized by the predominance of fallow lands. The age composition of crane flocks varied with acorn availability since more juveniles were reported in areas with relatively lower acorn availability. Time budget was also related to acorn availability, since cranes spent relatively more time preening in those wintering areas with higher availability of acorns. Globally evidence suggests the need to maintain the fallow land (posíos) to sustain the wintering populations of cranes in Iberia, since this agro-grazing system maintains most of the juvenile cranes using the western migratory route.     

Growth and yield of young Quercus ilex coppice stands in the Tafferte forest (Morocco)

The coppice method is the silvicultural system commonly applied to Quercus ilex (holm oak) forests for the production of firewood and charcoal. Although the holm oak is the species that covers the largest area in Morocco (1400 000 hectares), little is known about its growth and yield. The objective of this study is to quantify the stand growth and yield of young coppice holm oak stands in the Tafferete forest in the eastern part of the Middle Atlas (Morocco).Equations relating dominant height to age, and basal area, volume, and number of trees to dominant height were developed and used to derive young holm oak coppice yield estimates.The growth and yield information developed in this study may have a greater utility in defining cultural measures that should be applied in the early stage of holm-oak coppice stands.     

Genetic structure inside a declining red oak community in old-growth forest

Problems with oak regeneration have been documented in the last 50 years at numerous sites in the Midwestern United States. We applied nuclear microsatellites to examine the demographic and fine-scale spatial genetic structure of red oaks in two old-growth stands in Indiana. Oaks in one stand have declined in numbers over the past several decades whereas oaks in the other, smaller stand have increased. Large amounts of genetic variation were maintained within stands, and there was slight but significant differentiation among stands. There was significant but weak isolation by distance genetic structure within the large stand, likely reflecting family structure. No significant differences exist in allele frequencies or in levels of genetic diversity between cohorts that remain well represented within each stand, even between medium-sized adults and those antedating European settlement of the area. However, a virtual absence of smaller size classes in the forest interior of the large stand represents the early stages of a genetic bottleneck in what had been the core habitat of this stand. Whether future generations of this old-growth stand will retain the present genetic character depends on the oaks regenerating at the forest margins, absent any major changes in disturbance regimes. Similar demographic and genetic dynamics are likely occurring in a large number of remnant oak forests across the Midwest.     

Conifer regeneration failure in New Zealand: dynamics of montane Libocedrus bidwillii stands

Size and age structure analysis were used to reconstruct the population dynamics of six apparently old and stable stands dominated by the conifers Libocedrus bidwillii and/or Dacrydium biforme in Camp Creek, Westland, New Zealand. All stands contained relatively even-aged groups of Libocedrus that had probably originated after either mass movement or windthrow. The age distributions of stands reflected the effects of disturbances 200–400 yr ago.There was no evidence to support synchronous failure of regeneration of Libocedrus and Dacrydium biforme. Between 1600 and 1800 A.D., the period of the postulated conifer regeneration gap, over 70% of trees of these species had established. A lack of Libocedrus regeneration over the last 200 yr reflects a lack of disturbance over that time. Because of the dominating influence of disturbances and other factors it may be inappropriate to use tree population structures alone to substantiate or refute claims of recent climate change.     

Mediterranean pine and oak distribution in southern Spain: Is there a mismatch between regeneration and adult distribution?

Questions: What is the current distribution of pine and oak species along environmental gradients in southern Spain? Do pine and oak regeneration niches differ from the environmental niches of adults? Is oak species regeneration favoured under the canopy of pine forests? Location: Forest areas of Andalusia (～87 600 km², southern Spain). Methods: We compiled extensive forest inventory data to explore differences in abundance (basal area, m² ha^?1) patterns of adults (dbh >7.4 cm) and regeneration (dbh ≤7.4 cm) of five pine and five oak species. Canonical correspondence analysis (CCA) and generalized linear models were applied to explore species–environment relationships along climatic, edaphic, topographic and fire‐frequency gradients. Results: Both pines and oaks segregated along complex environmental gradients, with pines generally dominating in more severe (colder and drier) environments, while oaks dominated in milder, wetter winter areas. In 40‐55% of mature pine stands there was a lack of regeneration in the understorey, while in two oak species (Q. suber and Q. canariensis) 70% of stands did not show regeneration. Pine recruits were found at a higher frequency and abundance under the canopy of their congeners, whereas some oaks (Q. ilex) had greater regeneration under mixed pine–oak canopies. Conclusions: Climatic limitations and soil properties partly explained the regional distribution of pines and oaks. We found evidence for an upward shift of Q. ilex recruits towards areas with colder conditions in pine forests, which could be explained by a possible facilitative effect of the pine canopy on seedling establishment.     

Variation of stand structure and regeneration of Mediterranean holm oak along a grazing intensity gradient

Mediterranean ecosystems are traditionally influenced by human disturbance. In the mountain chains of south-eastern Spain, holm oak forests representing the potential natural vegetation are currently highly fragmented and subjected to extensive grazing by goats and sheep. This grazing activity by large herbivores has important impacts on community structure and dynamics of the remaining remnants because several processes associated with reproduction are affected. The present research analyses the variation of stand structure and regeneration of six forests dominated by Quercus ilex along a gradient of increasing grazing intensity. Stands of higher grazing intensity are characterized by a lower trunk number. Particularly, trees > 1.3 m height and diameter classes < 20 cm are clearly less represented. In addition, more intensively grazed forests show a highly aggregated dispersion of trees, which might be explained by an increase of clonally produced ramets by silvipastoral use. On the other hand, no Quercus ilex seedlings were found in any of the stands examined, although acorn production was high and did not correlate with grazing intensity. However, grazing clearly influences the quality of acorn harvest. We found a significant negative correlation between acorn length, acorn mass and grazing intensity and a positive correlation between the percentage of aborted acorns, acorn predation and herbivore activity. Thus, management of grazing in holm oak forests must take the high influence that livestock exerts on the regeneration of Quercus ilex into account, and maximum sustainable grazing intensity should be significantly lower than 0.8 animals per ha.     

Canopy seed bank structure in relation to: fire,tree size and density

To assess the canopy seed bank structure of Pinus halepensis, we measured the level of serotiny and the seed bank size and density of trees in unburned stands and post-fire regenerated stands in Israel. We analysed the effects of tree size, tree density and fire history on the level of serotiny. The level of serotiny decreased with an increase in tree height. The high level of serotiny in short trees could be explained by selection to increase regeneration chances after burning at pre-mature age. Also, limitation of long-distance seed dispersal opportunities in short trees may favour high serotiny levels. The level of serotiny was higher in post-fire stands than in unburned stands, suggesting a fast selection for serotiny by fire. Unburned stands had a higher total stand seed density than post-fire regenerated stands, but the proportion of seeds in serotinous cones of the total stand seed density was higher in post-fire regenerated stands. The fact that P. halepensis bears simultaneously serotinous and non-serotinous cones reflects its dual strategy as both a post-fire obligate seeder, mainly from serotinous cones and an early coloniser during fire-free periods, mainly from non-serotinous cones. The relative investment in these strategies is dependent on fire history and varies with tree height. Furthermore, mature brown cones can contribute to post-fire regeneration in case of spring fires, and serotinous cones are known to open partially also in dry spell events. Thus, post-fire regeneration and invasion are strategies, which seem to complement each other.     

Honeydew collection by the invasive garden ant Lasius neglectus versus the native ant L. grandis

Honeydew collection performed by the invasive ant Lasius neglectus and by the native ant L. grandis was compared. The invasive ant collected 2.09 kg of honeydew per tree while the native ant collected 0.82 kg. The aphid Lachnus roboris was visited by both ant species. In holm oaks colonized by L. neglectus, aphid abundance tended to increase and its honeydew production increased twofold. The percentage of untended aphids was lower in holm trees occupied by L. neglectus. As tending ants also prey on insects, we estimated the percentage of carried insects. The native ant workers carried more insects than the invasive ant. Both ant species preyed mainly on Psocoptera and the rarely tended aphid, Hoplocallis picta. We conclude that the higher honeydew collection achieved by L. neglectus was the consequence of (1) its greater abundance, which enabled this ant to tend more Lachnus roboris and (2) its greater level of attention towards promoting an increase of honeydew production. Handling editor: Heikki Hokkanen     

Spatial segregation of pines and oaks under different fire regimes in the Sierra Madre Occidental

Surface fire can modify spatial patterns and self-thinning in pine-oak ecosystems. Spatial pattern analyses were used to compare pattern development and interspecific spatial interactions in trees and seedlings in five Madrean pine-oak stands with different recent fire histories. Interspecific and intraspecific patterns were compared in small (< 15 cm dbh) and large (< 15 cm dbh) diameter classes of the pines (Pinus durangensis, P. teocote, and P. leiophylla) and oaks (Quercus sideroxylla, Q. crassifolia, and Q. laeta) that collectively dominated the five stands. Numbers of juvenile trees in 2.5 × 2.5 m subplots were correlated with cumulative distances to adult trees. Small pine and oak trees were intraspecifically clustered at all scales, irrespective of fire regime. Large pines were strongly clustered only in stands with longer fire-free intervals, and patterns of large versus small pine trees were regular or random in frequent fire stands. These patterns were consistent with fire-induced mortality of maturing trees under frequent fire. Large and small pines were segregated from small oaks at short and long distances in one stand with a 32-year fire-free interval, implying that two or more dynamic factors had produced regular patterns at different scales. Such regular spatial patterns at short distances were not seen in other stands. Therefore, there was little evidence for direct competition between oaks and pines. The results reported here are consistent with studies from other pine-oak ecosystems showing that different fire regime and site factors interact to influence stand development processes and relative dominance of pines and oaks. In some stands, the continued absence of fire could foster increasing tree densities and an intensification of local neighborhood effects, producing segregation of pine and oak species at longer distances. This revised version was published online in August 2006 with corrections to the Cover Date.     

影响不同林型天然红松混交林林隙更新的土壤特征因子

采用野外调查、样品采集和统计分析等相结合的方法,对小兴安岭天然红松混交林3种不同林型(椴树红松混交林(TP)、枫桦红松混交林(BP)、云冷杉红松混交林(PAP))的林隙及其邻近郁闭林分的土壤特征因子和树木更新的相关性进行了研究,旨在阐明林隙土壤特征因子对树木更新的影响,从而为小兴安岭天然红松混交林植被更新、退化生态系统的恢复和可持续经营提供基础数据和实践参考。结果表明:郁闭林分土壤有机质、全氮质量分数显著高于3种不同林型的林隙。有效磷和速效钾含量在BP内与其他林型之间差异显著。3种林型林隙内p H值均略高于其郁闭林分,但与其差异均不显著。3种林型林隙内更新总密度、幼树更新密度与郁闭林分差异显著(P0.05),PAP林隙中更新总密度和幼树更新密度最高。BP林隙面积与更新密度相关不显著,乔木幼苗、幼树更新密度与有机质(r=-0.400,r=-0.475)、全氮均呈显著负相关(r=-0.519,r=-0.603)。TP林隙内全氮与乔木幼苗更新密度呈正相关(r=0.092),而与乔木幼树更新密度呈显著负相关(r=-0.585)。PAP林隙内全氮与乔木幼苗、幼树更新密度均呈负相关。郁闭林分幼苗更新密度分别与有机质、全氮、速效钾、p H值、脲酶和蛋白酶呈负相关。主成分分析表明,全氮是影响林隙和郁闭林分树木更新的关键因素。     

Larval assemblages of large saproxylic cerambycids in Iberian oak forests: wood quality and host preference shape resource partitioning

Population Ecology - Oak open woodlands are widespread in southwestern Iberia and present outstanding ecological, biodiversity and socio-economic values. Evergreen species (holm and cork oaks) are...     

Are sub-alpine firs evolving towards semelparity?

Summary In sub-alpine forests of Northeast America and Japan pure stands of trees in the genusAbies exhibit wave regeneration. Opportunities for recruitment in such forests are confined to a window in time and space that coincides with the death of an even-aged cohort of adult trees. I suggest that the coincidence of this recruitment window with the death of adults at a predictable age should select for convergence between age at first reproduction and age at death. Ultimately this would lead to the evolution of semelparity. The available evidence supports this hypothesis for wave-regeneratedAbies populations in Japan. A field test of the hypothesis is also suggested.     

Stand structure and dynamics of wave-typeAbies sachalinensis coastal forest

Stand structure and regeneration pattern were examined inAbies sachalinensis coastal forest in northern Hokkaido. In the forest a similar phenomenon to the wave regeneration in subalpine forests has been found. Wave regeneration has been reported for montaneAbies forests in central Japan and North America. Differences and similarities between wave-type stands in this coastal forest and wave-regenerated montane forests were clarified. The shift of dead tree zone, stand structure and regeneration pattern in wave-type stands are the same as in subalpine wave-regenerated forests. High density of individuals is considered to be an internal factor which causes stand-level dieback and also enables the stands to persist in the severe environment in both forests. A difference between wave-regenerated forests andA. sachalinensis wave-type stands is the number of dead tree zones, which is only one in wave-type stands. Changes of regeneration patterns ofAbies sachalinensis with environmental gradient from seaward to inland were related to this difference.     

Variations in levels of clonality among Populus nigra L. stands of different ages

The evolution of genotypic diversity with population age remains poorly explored in clonal plant populations despite the potential for important shifts to occur through the course of time. Woody sprouting species are particularly under-represented in studies investigating intra-specific variations in levels of clonality from one locality to the next and through time. In this study we sought to determine the incidence and frequency of replicate genotypes in natural Populus nigra L. (Salicaceae) stands of different ages. Ten stands of this woody riparian sprouting species were selected in each of three distinct age groups (young, middle-aged and old) along a 30 km stretch of the River Garonne (south-west France). Leaf samples were collected from 15 neighbouring trees in each stand (450 samples in total) and replicate genotypes were identified using five SSR markers. Replicate genotypes were identified in two-thirds of all stands sampled (i.e. 50 of young stands, 100 of middle-aged stands and 50% of old stands). Young stands had significantly fewer replicated genotypes than middle-aged or old stands, while middle-aged stands had the greatest number of replicated genotypes. Replicate genotypes were most often found to occur as nearest neighbours and formed relatively small, discrete units (i.e. 2–4 trees growing in close proximity to one another). This suggests that asexual regeneration frequently occurs through flood-training in this species, although asexual regeneration from translocated fragments also evidently occurs as evidenced by 11 cases of replicate genotypes occurring in widely separated stands (up to 19 km apart). The results of this study highlight the need for a hierarchical sampling strategy in space and across age groups for an accureate understanding of the genotypic structure of woody sprouting species populations. Conservation and management of effective population sizes will benefit from better insight into not only spatial, but also temporal variations in levels of genotypic diversity.Co-ordinating editor: J. Tuomi