Oak trees and soil interactions in Mediterranean forests: a positive feedback model

Questions: What is the spectrum of variability of chemical elements in a Mediterranean forest ecosystem across the different compartments? Do co‐existing tree species with different leaf chemical composition and nutrient cycling distinctly modify soil conditions? Could these species‐specific, tree‐generated soil changes create a potential positive feedback by affecting long‐term species distribution? Location: Mixed oak forests of southern Spain, Los Alcornocales Natural Park. Methods: We sampled and chemically analysed five different ecosystem components: leaves, leaf fall, litter and superficial (0–25 cm) and sub‐superficial (25–50 cm) soil beneath the canopies of evergreen Quercus suber and deciduous Q. canariensis trees. We used multiple co‐inertia analysis (MCoA) to conjointly analyse the patterns of variability and covariation of eight macro‐ and micronutrients determined in each of the sampled ecological materials. We implemented a path analysis to investigate alternative causal models of relationships among the chemical properties of the different ecosystem components. Results: Variability in the concentration of chemical elements was related to the nature of their biogeochemical cycles. However, the rank of element concentration was consistent across ecosystem components. Analysis of co‐inertia (MCoA) revealed that there was a common underlying multivariate pattern of nutrient enrichment in the ecosystem, which supported the hypothesis of a separation in biogeochemical niches between the two co‐existing oak species, with Q. canariensis having richer plant tissues and more fertile soil directly under each tree than Q. suber. The feasibility of a potential tree–soil positive feedback model was the only statistically validated among several alternative (non‐feedback) models tested. Conclusions: In the studied Mediterranean forests, oak species distinctly modify soil fertility conditions through different nutrient return pathways. Further investigation is needed to address whether these tree‐generated soil changes could affect seedling establishment and ultimately influence species distribution.     

Coexistent Mediterranean woody species as a driving factor of Phytophthora cinnamomi infectivity and survival

The long-term conservation of Mediterranean mixed oak forests is seriously threatened by the massive mortality of Quercus suber caused by the exotic pathogen Phytophthora cinnamomi. This species frequently grows in mixed forests under natural conditions, but nothing is known about how its level of disease might be altered by the diversity and identity of coexisting neighbours varying in susceptibility to the exotic pathogen. Here we analysed the individual and combined effects of Q. suber and the main coexisting tree species (Quercus canariensis and Olea europaea subsp. europaea var. sylvestris) in mixed forests of southern Spain on the production of infective and survival spores of P. cinnamomi. Through in vitro experiments, it was demonstrated that mixtures of Q. suber and Q. canariensis highly stimulated the production of P. cinnamomi zoospores in comparison with both species in monocultures. Olea europaea did not stimulate zoospore production. Under controlled conditions, the initial and final densities of inoculum in soil planted with monocultures of O. europaea and Q. canariensis did not differ. However, inoculum densities significantly decreased along the experiment in Q. suber mixtures with O. europaea and Q. canariensis. Phytophthora cinnamomi was able to infect and cause root rot symptoms on all tree species, including O. europaea var. sylvestris. We concluded that mixed stands of Q. suber and Q. canariensis are able to stimulate P. cinnamomi infectivity and survival much more than monospecific stands, and consequently under favourable conditions for root disease development, the coexistence of Q. suber and Q. canariensis might exacerbate Mediterranean forests decline. This study also constitutes the first report of O. europaea var. sylvestris as host and inductor of P. cinnamomi sporulation under controlled conditions.     

Seed removal in two coexisting oak species: ecological consequences of seed size, plant cover and seed-drop timing

Seed predation and dispersal can critically influence plant community structure and dynamics. Inter‐specific differences arising at these early stages play a crucial role on tree recruitment patterns, which in turn could influence forest dynamics and species segregation in heterogeneous environments such as Mediterranean forests. We investigated removal rates from acorns set onto the ground in two coexisting Mediterranean oak species –Quercus canariensis and Q. suber– in southern Spain. We developed maximum likelihood estimators to investigate the main factors controlling probabilities of seed removal and to describe species‐specific functional responses. To account for inter‐specific differences in seed‐drop timing, two experiments were established: a simultaneous exposure of acorns of the two species (synchronous experiments) and a seed exposure following their natural seed‐drop phenology (diachronic experiments). A total of 1536 acorns were experimentally distributed along a wide and natural gradient of plant cover, and removal was periodically monitored for three months at two consecutive years (with contrasting differences in seed production and thus seed availability on the ground). The probability of seed removal increased with plant cover (leaf area index, LAI) for the two oak species. Inter‐specific differences in acorn removal were higher in open areas and disappeared in closed microhabitats, especially during a non‐mast year. Despite later seed‐drop, Q. suber acorns were removed faster and at a higher proportion than those of Q. canariensis. The higher probability of seed removal for this species could be attributed to its larger seed size compared to Q. canariensis, as inter‐specific differences were less pronounced when similar sized acorns were exposed. Inter‐specific differences in seed removal, arising from seed size variability and microsite heterogeneity, could be of paramount importance in oak species niche separation, driving stand dynamics and composition along environmental gradients.     

Proteomic analysis from haploid and diploid embryos of Quercus suber L. identifies qualitative and quantitative differential expression patterns

Quercus suber L. is a Mediterranean forest species with ecological, social and economic value. Clonal propagation of Q. suber elite trees has been successfully obtained from in vitro‐derived somatic and gametic embryos. These clonal lines play a main role in breeding and genetic studies of Q. suber. To aid in unravelling diverse genetic and biological unknowns, a proteomic approach is proposed. The proteomic analysis of Q. suber somatic and gametic in vitro culture‐derived embryos, based on DIGE and MALDI‐MS, has produced for the first time proteomic data on this species. Seventeen differentially expressed proteins have been identified which display significantly altered levels between gametic and somatic embryos. These proteins are involved in a variety of cellular processes, most of which had been neither previously associated with embryo development nor identified in the genus Quercus. Some of these proteins are involved in stress and pollen development and others play a role in the metabolism of tannins and phenylpropanoids, which represent two of the major pathways for the synthesis of cork chemical components. Furthermore, the augmented expression levels found for specific proteins are probably related to the homozygous state of a doubled‐haploid sample. Proteins involved in synthesis of cork components can be detected at such early stages of development, showing the potential of the method to be useful in searching for biomarkers related to cork quality.     

Rodent acorn selection in a Mediterranean oak landscape

Quercus suber, Quercus ilex and Quercus coccifera (Cork, Holm and Kermes oaks, respectively) are common evergreen oak species that coexist in the landscapes of the western part of the Mediterranean basin. Rodents are the main acorn predators and thus one of the main factors for understanding recruitment patterns in oaks. In this paper we analyse to what extent mice prefer acorns from one oak species over another in three oak species studied using acorn removal experiments and video tape recordings. Twenty labelled acorns from each of the three Quercus species (60 acorns) were placed in 40 cm×40 cm quadrats on each plot. Because selection might vary as a result of the vegetation context, we performed the trials in the five main vegetation types within the study area (four replicates in each vegetation type) in order to control for habitat influences on rodent acorn preferences (a total of 20 plots). The removal of 1,200 acorns occurred within 68 days. Mice removed 98.7% of the acorns. Q. ilex acorns were preferred over Q. suber and Q. coccifera in all vegetation types except in pine forest, where no acorn preferences were detected. Acorn removal rates differed with vegetation type, correlating positively with shrub cover. The distance at which acorns were displaced by rodents (mean =4.6 m±5.1 SD) did not differ between acorn species, but varied among vegetation types. Bigger acorns of Q. coccifera were selected only after Q. ilex and Q. suber acorns were depleted, while no size selection was detected for the latter two species. Thus, we conclude that rodents show preference for some oak acorns and that landscape context contributes significantly to rodent activities and decisions.     

Not only size matters: Acorn selection by the European jay (Garrulus glandarius)

A strong selection for acorn characteristics is expected to have evolved in the mutualistic relationship between the European jay (Garrulus glandarius) and the oak (Quercus spp.). Bossema's pioneer work suggested that jays do not select acorns randomly, but rather they preferentially select some size and species. Preference for some seeds over others may have implications on plant community dynamics by conferring advantages (or disadvantages) on the selected (avoided) seed characteristics. In this paper we test to what extent jays select acorns by species and/or by size and the relation between these two traits in Mediterranean oak species. The experiments consist of a set of field tests in which acorns from four different coexisting Mediterranean oak species (Quercus ilex, Quercus faginea, Quercus suber, and Quercus coccifera) were placed in artificial feeders accessible to wild jays. The acorns were previously measured to control individual acorn characteristics. Using video-recording techniques, we followed jay activity and the fate of each acorn (sequence of acorn selection and method of transport). Q. ilex acorns were preferred over other acorns, and Q. coccifera acorns were avoided when other acorns were available. Preference for Q. faginea and Q. suber acorns was intermediate, that is, they were preferred over Q. coccifera acorns but not over Q. ilex acorns. Large acorns were also preferred although acorn species selection was stronger than size selection. Jays selected species and size both by visual means and by using acorn area as an indicator of size. Acorns wider than 17–19 mm were carried in the bill because of throat limitation. Our results confirm Bossema's study on temperate oaks and extend it to Mediterranean oak species, revealing implications on mixed oak forest dynamics.     

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.     

Patterns and ecological consequences of abiotic heterogeneity in managed cork oak forests of Southern Spain

Spatial heterogeneity of abiotic factors influences the structure and function of forests and must be taken into account for their conservation and sustainable management. In this study, we evaluate the heterogeneity of abiotic environmental variables in managed cork oak (Quercus suber L.) forests in southern Spain at patch, site and regional scales. The extent of spatial heterogeneity depended on the environmental variable examined and the scale considered. For example, soil Mn and P and light availability in the understorey were very heterogeneous at the regional scale, while soil N had low regional heterogeneity, but high spatial variability, at patch scale, attributed to open overstorey and grazing disturbance. There was a general trend of increasing heterogeneity with spatial scale. We also study the effects of a silvicultural practice—shrub clearing on the forest environment and its consequence for spatial heterogeneity. Shrub clearing increased understorey light and decreased its spatial heterogeneity with idiosyncratic effects on soil properties and their spatial heterogeneity at each site. Finally, we compare the heterogeneity (estimated by the coefficient of variation) obtained in these cork oak forests with a database compiled from published studies on other forest environments. The comparison revealed a remarkable extent of abiotic heterogeneity in the cork oak forests studied, suggesting that a sustainable management of these forests should combine intrinsic and human induced abiotic heterogeneity to preserve crucial ecological processes and to maintain high levels of biodiversity.     

Allozyme variation in cork oak (Quercus suber L.): the role of phylogeography and genetic introgression by other Mediterranean oak species and human activities

 Genetic variation in the cork oak (Quercus suber L.) was investigated using 11 loci from seven enzyme systems in 40 populations sampled over the entire distribution of this species in the western Mediterranean Basin. Mean heterozygosity values over the polymorphic loci (Ho=0.283), the percentage of polymorphic populations (M=0.76), and the total genetic diversity (Ht=0.31) from which 11% was accounted for among-population variation, are among the highest recorded in oak species. In contrast to previous results in holm oak (Q. ilex L.), another evergreen species in the same area, cork oak possessed a smaller allele pool and a lower average number of alleles per locus and per population (A=2.0). More particularly, very few low-frequency alleles were observed in cork oak except for eight populations in which allozyme polymorphism at locus Pgi 1, diagnostic between both species, indicates that these low-frequency alleles are introgressed from holm oak. On the basis of the genetic distance estimated from allozyme frequencies, 32 of the 40 cork oak populations studied were classified into two very distinct sets which also corresponded to distinct geographic areas. One set gathered together the 18 populations from the Iberian peninsula and two adjacent areas in France, i.e. the centre of origin of cork oak, according to paleobotanical data. This set was characterized by a larger allele pool, a higher within-population genetic diversity and a lower differentiation between populations than was observed in the other set, which comprised the populations from North Africa, Sicily, Sardinia, Corsica, continental Italy and the region of Provence (southeastern France). In these more southern and eastern disjunct areas, cork oak migration from Iberia may have occurred at different periods since the end of the Tertiary. The possible effect of human activity on cork oak genetic structure, i.e. the selection of good-quality cork, acorn over-use for animal food, and even human nutrition, is discussed. Received: 3 March 1998 / Accepted: 19 March 1998     

The Balearic Islands: a reservoir of cpDNA genetic variation for evergreen oaks

Aim To analyse the role of the Balearic Islands as a refuge area for evergreen Quercus (cork oak: Quercus suber L., holm oak: Q. ilex L., kermes oak: Q. coccifera L.), by using molecular, historical and palaeobotanical data. Location The Western Mediterranean Basin (Balearic Islands, eastern Iberia, Provence, Sardinia, Corsica, Sicily, Malta, Italy, Northern Africa). Methods We sampled 108 populations and used the PCR‐RFLP technique with five universal cpDNA primers to define haplotypes in the sampled populations. Diversity, differentiation parameters and spatial analysis of the populations, using a spatial version of amova , were linked to the geological history of the Western Mediterranean Basin in order to explain the present spatial pattern of the evergreen Quercus populations in the Balearics. Results Evergreen Quercus cpDNA shows a complex structure, with remnants of ancient diversity in the Balearics. Balearic populations of holm oak are related to Iberian populations, while for cork and kermes oaks, we found both Tyrrhenian and Iberian haplotypes. Main conclusions The complex spatial patterns of cpDNA in Balearic evergreen Quercus appears explicable in terms of a combination of physical (vicariance and long distance dispersal) and biological (introgressive hybridization) factors. The Balearics constitute a glacial refuge area and a reservoir of genetic variation with traces of ancient diversity from Messinian–Pliocene stages.     

Tree species identity and forest composition affect the number of oak processionary moth captured in pheromone traps and the intensity of larval defoliation

1. Pure forests are often seen as being more prone to damage by specialist pest insects than mixed forests, and particularly mixed forests associating host and nonhost species. We addressed the effect of tree diversity on oak colonization and defoliation by a major specialist pest, the oak processionary moth (OPM)
2. We quantified the number of male OPM moths captured and larval defoliation in pure stands of two oak host species (Quercus robur and Quercus petraea) and in mixed stands associating the two oak species or each oak species with another nonhost broadleaved species. We conducted two complementary studies to test the effect of host species and stand composition: (i) we used pheromone trapping to compare the number of males OPM captured throughout the distribution of oak hosts in France and (ii) we noted the presence of OPM nests and estimated defoliation in mature forests of north‐eastern France.
3. Oak species and stand composition significantly influenced the number of male OPM captured and defoliation by OPM larvae. Quercus petraea was consistently more attractive to and more defoliated by OPM than Q. robur. Both oak trees were attacked more in pure stands than in mixed stands, in particular mixed stands associating oaks with another (nonhost) broadleaved species.
4. The results of the present study support the view that mixed forests are more resistant to specialist pest insects than pure stands, and also indicate that this trend depends on forest composition. Our study provides new insights into OPM ecology and has potential implications for forest management, including the management of urban forests where OPM causes serious human health issues.

     

Regeneration of a marginal Quercus suber forest in the eastern Iberian Peninsula

Question: Small and marginal forest populations are a focus of attention because of their high biodiversity value as well as the risk of population decline and loss. In this context, we ask to what extent a small, marginal Quercus suber (Cork oak) population located in the eastern Iberian Peninsula (Valencia, Spain) has the capacity for self‐regeneration and what are the factors that determine its recruitment variability. Location: Quercus suber forest in Pinet (Valencia, Spain). Methods: We performed a spatially explicit sampling both of the recruitment and of the potential parameters that could account for the recruitment variability. Using regression techniques we model the recruitment occurrence and abundance, and then we test to what extent the model obtained is still constrained by the spatial dependence. Results: Quercus suber recruitment density ranges from 0 to 18.66 individuals/25m² (mean = 1.46, SD = 2.8), with a very skewed distribution. Recruitment is similar under Q. suber forests and under Pinus forests, but it is almost absent under shrublands. Thus the parameters that explain most of the recruitment variability in local vegetation types are: the presence and cover of shrubs (negative relationship with recruitment), the basal area of Q. suber and Pinus and the amount of bare soil (all positively related to recruitment). These parameters are strongly related to the ecological processes driving recruitment (i.e. dispersal and predation) and they remove most of the spatial dependence of recruitment. Most recruiters, however, are small, forming a seedling bank rather than growing to successfully colonize new habitats. Conclusion: The results suggest that although recruitment densities are not very high, they do not limit potential regeneration in the Pinet Q. suber forest. However, successful regeneration is not observed. If we aim to increase the Pinet Q. suber population size, land management measures need to provide appropriate conditions for both seedling establishment in shrublands (e.g. shrub clearing) and seedling growth in woodlands (e.g. Pinus logging).     

Multiple Recruitment Limitation Causes Arrested Succession in Mediterranean Cork Oak Systems

Lack of tree regeneration and persistency of species-poor shrublands represent a growing problem across Mediterranean evergreen oak forests. What constrains forest regeneration is poorly understood, and restoration attempts have been largely unsuccessful. We assessed the contribution of four different mechanisms of tree recruitment limitation (that is, source, dispersal, germination, and establishment) in a cork oak (Quercus suber) system in southern Portugal. Using a combination of field studies and experiments, we quantified seed production, seed removal and dispersal, seed survival and germination, seedling establishment and survival, as well as cork oak natural regeneration for the three dominant vegetation types in this system (Cistus ladanifer shrubland, oak forest, and oak savanna). We found that all four forms of cork oak recruitment limitation were significantly more severe in shrublands than in oak forests and savannas, so that oak seedling recruitment in shrubland was impeded in multiple ways. Our results explain why transitions from shrublands to oak savannas and forests are extremely difficult, and that the release from arrested succession in this system requires the simultaneous relief of multiple constraints on recruitment limitation in the early life history of oaks. These results have important implications for the restoration and conservation of Mediterranean oak systems.     

Ecological and biogeographical analyses of Mediterranean forests of northern Morocco

Mediterranean forests in northern Morocco have been studied. Tree species composition and abundance were investigated in 84 forest sites, distributed throughout the sandstone formation of the Tangier region. The relative abundance of 15 tree and arborescent shrub species was related, by ordination analysis, to altitude, disturbance by fire, and soil fertility. High-elevation forests were dominated by Cedrus atlantica, Quercus pyrenaica and Pinus pinaster. Sacred forests (protected as holy burial places) had a distinct stand structure, and were considered as refuges, for maintaining biodiversity at landscape scale. The evergreen oak Quercus suber and the semideciduous Quercus canariensis co-dominated the lowland forests. The mean basal area of the studied forests was 34 m² ha^–1. The mean trunk size (dbh) for Q. suber was 24 cm and for Q. canariensis 36.5 cm; in both cases there were indications of declining regeneration at population level. A comparative biogeographical analysis with the equivalent region in southern Spain (separated by the Strait of Gibraltar; 14 km wide), indicated a possible biodiversity loss in the Moroccan forests.     

Acorn preference under field and laboratory conditions by two flightless Iberian dung beetle species (Thorectes baraudi and Jekelius nitidus): implications for recruitment and management of oak forests in central Spain

1. Recently, a mutualistic relationship has been described between some dung beetles (Thorectes lusitanicus and Mycotrupes lethroides) and oak species (Quercus suber, Q. canariensis, and Q. rubra), which could be crucial for ensuring seedling recruitment and sustaining the equilibrium of oak populations. For T. lusitanicus, a diet based on acorns during the reproductive period improved resistance to low‐temperature conditions and improved ovarian development. 2. In this paper, we conducted field and laboratory experiments to investigate the interaction between two potential acorn‐eating beetles, Thorectes baraudi and Jekelius nitidus, with Quercus suber. We determined the feeding preferences of both beetle species and estimated the rates of acorn manipulation by beetles according to habitat structure and several characteristics of the acorn, such as seed size and acorn infestation by weevils. 3. Results demonstrated the positive interaction between the dung beetle Thorectes baraudi and Quercus trees. Thorectes baraudi was clearly more attracted to volatiles of acorns than to dung. Jekelius nitidus, on the contrary, was either not or anecdotally attracted to acorns. On the contrary, in the case of Jekelius nitidus, the acorn attraction could be considered anecdotal or even accidental. Our field results demonstrated the acorn burying behaviour of T. baraudi in the oak forests of the Cabañeros National Park (Spain), suggesting a potential role of this beetle species as an active secondary acorn disperser. 4. This unexpected behaviour could be particularly important in Mediterranean oak forests and savannahs, where most Quercus species are strongly recruitment limited because of serious overgrazing problems.     

Field trapping of the flathead oak borer Coroebus undatus (Coleoptera: Buprestidae) with different traps and volatile lures

The flathead oak borer Coroebus undatus F. (Coleoptera: Buprestidae) is one of the primary pests of cork oak Quercus suber L. in the Mediterranean region causing great economic losses to the cork industry. Very little is known about its biology and behavior and, so far, no control measures have been established. We present the results of a pilot study aimed to develop an efficient trapping method for monitoring this harmful pest. In a 3‐year field study, purple‐colored prism traps baited with a mixture of green leaf volatiles (GLVs) from the host have been shown the most effective combination to catch C. undatus adults (solely females) compared to other trap and lure types tested. Wavelength and reflectance measurements revealed that purple traps exhibit reflectance peak values similar to those found in the abdominal and elytral cuticle of both sexes, suggesting the involvement of visual cues for mate location in this species. The data presented are the first to demonstrate captures of adults of the genus Coroebus by an attractant‐based trapping method.     

Differential ecophysiological responses to seasonal drought of three co-existing oak species in northern Greece

Oaks (Quercus spp.) represent the most important broadleaf genus with respect to forest-shaping tree species in the Mediterranean. Considering future climate scenarios (increased drought conditions), the identification of drought tolerant oak species is of great importance for future forest management in this region. The objective of the study was the comparison of physiological status of three economically and ecologically valuable oak species (Quercus ilex, Quercus frainetto and Quercus pubescens) co-existing in natural coppice stands in NE Greece, in response to seasonal drought stress. Measurements were conducted between June and September 2016, every 15–20 days until leaf falling. The parameters studied were predawn leaf water potential and fast chlorophyll fluorescence induction curves (OJIP test), chlorophyll content, and relative water content. Meteorological data from the area were also collected. Photosynthetic parameters such as performance indices (PI_abs and PI_tot) reacted to summer drought conditions, with Q. frainetto showing the lowest values. The discrepancy between species increased with duration of drought period. Q. frainetto revealed the lowest predawn water potential values. The results indicate that Q. frainetto is less suitable for future forestry applications in the studied climate/elevation zone than Q. pubescens and Q. ilex.     

A comprehensive assessment of the transcriptome of cork oak (Quercus suber) through EST sequencing

Background

Cork oak (Quercus suber) is one of the rare trees with the ability to produce cork, a material widely used to make wine bottle stoppers, flooring and insulation materials, among many other uses. The molecular mechanisms of cork formation are still poorly understood, in great part due to the difficulty in studying a species with a long life-cycle and for which there is scarce molecular/genomic information. Cork oak forests are of great ecological importance and represent a major economic and social resource in Southern Europe and Northern Africa. However, global warming is threatening the cork oak forests by imposing thermal, hydric and many types of novel biotic stresses. Despite the economic and social value of the Q. suber species, few genomic resources have been developed, useful for biotechnological applications and improved forest management.

Results

We generated in excess of 7 million sequence reads, by pyrosequencing 21 normalized cDNA libraries derived from multiple Q. suber tissues and organs, developmental stages and physiological conditions. We deployed a stringent sequence processing and assembly pipeline that resulted in the identification of ~159,000 unigenes. These were annotated according to their similarity to known plant genes, to known Interpro domains, GO classes and E.C. numbers. The phylogenetic extent of this ESTs set was investigated, and we found that cork oak revealed a significant new gene space that is not covered by other model species or EST sequencing projects. The raw data, as well as the full annotated assembly, are now available to the community in a dedicated web portal at http://www.corkoakdb.org.

Conclusions

This genomic resource represents the first trancriptome study in a cork producing species. It can be explored to develop new tools and approaches to understand stress responses and developmental processes in forest trees, as well as the molecular cascades underlying cork differentiation and disease response.     

Molecular phylogeny and genetic diversity of Tunisian Quercus species using chloroplast DNA CAPS markers

Chloroplast DNA variation was studied in five evergreen Quercus species from the Western part of Tunisia using Cleaved Amplified Polymorphic Sequence (CAPS) technique. Five primer pair/endonuclease combinations have been used. Chlorotypes of Quercus species have been identified. The enzyme HinfI was more efficient in detecting polymorphism in oak species than TaqI. The phenogram showed five groups defining the five studied oak species: suber group, afares group, coccifera group, canariensis group and ilex group. The topology of phenogram showed that the classification depends only on species and independently of their geographic origin. The principal component analysis (ACP) corroborated the results of the tree branching and confirmed the existence of five species groups. Our results showed a genetic proximity between Quercus afares and Quercus coccifera species that may be due to temperature tolerance or the demographic history of these species. Nevertheless, a high value of G_ST calculated (G_ST = 1), suggesting that the maximum of variation is maintained among oak species. This result was confirmed by the low value of the genetic diversity within species (h_S = 0), the value obtained of the total genetic diversity (h_T = 0.378) and the absence of gene flow between species (N_m = 0). A high genetic proximity has been registered between Q. afares, Quercus suber and Quercus canariensis. Moreover, Q. afares shared the chlorotype of Q. suber and Q. canariensis which suggests its hybrid origin.     

Plant diversity patterns in the Aljibe Mountains (S. Spain): a comprehensive account

The Aljibe Mountains are located in the southern tip of the Iberian Peninsula and have a remarkable biogeographical interest. The complete plant species list (trees, climbers, shrubs, perennial and annual herbs, ferns, lichens, bryophytes and macroscopic algae) was recorded in four 0.1 ha plots from each of the most representative community types (Quercus suber woodland, Q. canariensis forest, open heathland and Q. coccifera shrubland). Up to 119 plant species were found in total in the Q. suber woodland plot. The diversity of woody plants was analysed from 44 samples of cover (100 m line), and the herbaceous layer was explored in 200 quadrats (of 0.5 × 0.5 m). Three biodiversity components (species richness, endemism, and taxonomic singularity) were evaluated in both shrub and herbaceous layers. Open heathlands showed the highest richness of endemic species, both woody and herbaceous. The highest number of woody species was found in the evergreen Q. suber woodland, and of herbaceous species in the semi-deciduous Q. canariensis woodland. Taxonomic singularity was higher in Q. canariensis woodlands and Q. coccifera shrublands for woody species, but there were no significant differences in the herbaceous layer. Local species diversity of heathlands in this region resembles that of South African heathlands (fynbos), despite the obvious geographic and floristic distance, and contrasts with the low diversity of biogeographically closer, European temperate heathlands. The Aljibe Mountains show high diversity values for different life forms (from trees to mosses) and spatial scales (from community to region), and are rich in endemic species. Thus, this area should be recognised as a relevant unit within the Mediterranean plant diversity hot spots.