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.     

Acorn dispersal estimated by radio-tracking

Bird-dispersed seeds are difficult to track, especially in the case of long-distance dispersal events. To estimate the oak dispersal distance and the seed shadow generated by the European jay (Garrulus glandarius), we inserted radio-transmitters in 239 acorns, placed them in bird-feeders and then located them by radio-tracking. Using this methodology we located the exact caching site of 94 Quercus ilex and 54 Q. suber acorns and determined the caching habitat characteristics (vegetation type, distance, spatial distribution). The results show that: (1) there is no differences in the dispersal distance distribution between the different acorn species or sizes, (2) dispersal distances range from approximately 3 m up to approximately 550 m (mean = 68.6 m; median = 49.2 m), (3) recently abandoned fields and forest tracks were the sites preferred by jays to cache acorns, whereas fields and shrublands were avoided and (4) seed shadows showed acorn aggregation zones (i.e. clusters of caches) close to the feeder as well as isolated caches at longer distances. The results also suggest that radio-transmitters are a cheap and reliable way to determine seed shadows and quantify both seed dispersal and post-dispersal seed predation for medium to large seeds.     

Acorn selection by Mexican jays: a test of a tri-trophic symbiotic relationship hypothesis

By caching acorns, jays serve as important dispersal agents for oak (Quercus) species. Yet little is known about which acorn characteristics affect selection by jays. In the traditional model of jay/oak symbiosis, large, brown, ripe acorns free of invertebrate parasites (e.g., Curculio acorn weevils) are selected by jays. Recently, it has been suggested that a tri-trophic relationship between oaks, jays, and weevils may have evolved to counter the negative dietary effects of acorn tannins. Under the tri-trophic model, jays would preferentially select acorns containing weevil larvae. We tested the assumptions that (1) acorns containing curculionid larvae exist in sufficient quantities to support jay populations and (2) jays can detect, and preferentially select, acorns containing weevil larvae, and investigated the cues by which jays select acorns. Captive Mexican jays (Aphelocomaultramarina) were presented Emory oak (Quercusemoryi) acorns in aviary feeding trials. Large, dense, viable acorns free of curculionid larvae were preferentially selected. Contrary to results of previous research, color did not affect selection. Acorn viability increased and curculionid larval occupancy decreased in adjacent savannas and isolated stands relative to existing oak woodland, perhaps favoring oak recruitment into adjacent lower-elevation grasslands. Our results compel us to reject the tri-trophic model for this system, and are consistent with the traditional jay/oak symbiosis model. Relatively long-distance dispersal of viable acorns favors Emory oak replacement, and spatial patterns of acorn viability and curculionid parasitism suggest expansion of Emory oak into adjacent low-elevation semi-arid grasslands. Received: 29 February 1996 / Accepted: 26 September 1996     

Summer-drought constrains the phenology and growth of two coexisting Mediterranean oaks with contrasting leaf habit: implications for their persistence and reproduction

This study analyses how coexisting evergreen and deciduous oaks adjust their phenology to cope with the stressful Mediterranean summer conditions. We test the hypothesis that the vegetative and reproductive growth of the winter deciduous (Quercus faginea Lam.) is more affected by summer drought than that of the evergreen [Quercus ilex L. subsp. ballota (Desf.) Samp.]. First, we assessed the complete aboveground phenology of both species during two consecutive years. Shoot and litter production and bud, acorn and secondary growth were monitored monthly. Second, we identified several parameters affected by summer conditions: apical bud size, individual leaf area (LA), leaf mass per area (LMA) and acorn yield in both species, and leaf-fall in Q. faginea; and analysed their variation over 10 years. Q. ilex performed up to 25% of shoot growth and most leaf development during summer, whereas Q. faginea completed most of both phenophases during spring. Secondary growth was arrested in summer under drought conditions. Approximately, 30–40% of bud and 40–50% of acorn growth was undertaken during summer in both species. Summer drought related to differences in LA, LMA and leaf senescence, but not to acorn yield. Both species had similar year-to-year patterns of acorn production, though yields were always lower in Q. faginea. Bud size decreased severely in both species during extremely dry years. In Q. ilex, bud size tended to alternate between years of large and small buds, and these patterns were followed by opposite trends in stem length. In Q. faginea, bud size was more stable through time. Q. ilex was more phenologically active during summer than Q. faginea, indicating a higher tolerance to drought. Furthermore, bud and fruit growth (the only two phenophases that both species performed during summer) were more severely affected by summer drought in Q. faginea than in the evergreen. The differential effects of summer drought on key phenophases for the persistence (bud growth) and colonization ability (fruit production) of both species may have consequences for their coexistence.     

Ribosomal RNA genes inQuercus spp. (Fagaceae)

The taxonomy of the genusQuercus is still unclear. In order to elucidate the taxonomy of Mediterranean oaks we have analyzed ribosomal RNA genes ofQuercus cerris, Q. coccifera, Q. trojana, Q. ilex, Q. suber, andQ. macrolepis by means of Southern blot hybridization. Oak nuclear DNA was extracted from root tips of 300 acorns and from catkins of single plants. EcoRI and BamHI restriction endonucleases were used. DNA electrophoresis and rRNA/DNA hybridization were performed usingVicia faba rRNA 18 S and 25 S as probes. The rRNA genes of all the species studied have an identical restriction mapping in the 18 S and 25 S regions, while differences in length are present in the intergenic regions.Q. cerris possesses at least four types of genes of 12.1, 11.5, 8.5, and 8.3 kb;Q. coccifera at least three types of 12.4, 10.4, and 10.1 kb;Q. trojana possesses the same rRNA genes asQ. cerris plus another gene type 12.0 kb long, with EcoRI and BamHI restriction sites in the intergenic spacer;Q. ilex at least three types of 12.4, 10.85, and 9.5 kb;Q. suber at least five types of 11.5, 11.0, 8.6, 8.5, and 8.3 kb;Q. macrolepis, finally, at least seven types of 11.5, 11.0, 10.2, 8.6, 8.5, 8.3, and 8.15 kb.Q. coccifera andQ. ilex rDNA appears quite different respect to other species examined, while high similarity seems to exist betweenQ. cerris, Q. trojana, Q. suber, andQ. macrolepis. These results are in agreement with the taxonomic model proposed bySchwarz for the genusQuercus.     

Effects of moderate shade and irrigation with eutrophicated water on the nitrogen economy of Mediterranean oak seedlings

We evaluated the effects of moderate shade (43% vs. 100% of full sunlight) and irrigation with eutrophicated river water (daily vs. alternate-day watering) on growth and nitrogen economy of seedlings of three Mediterranean oak species, two evergreen (Quercus coccifera, Quercus ilex subsp. ballota) and a deciduous (Quercus faginea), grown in pots outdoors. Seedling biomass, N pool, N concentration and N losses by litter fall were measured at the beginning (March 2002) and end (November 2002) of a growing season. All species showed an increase of biomass and N pool under shade and/or high irrigation, while only Q. coccifera – from more arid regions – did the same under full sunlight and low irrigation. At the end of the experiment, biomass of the evergreens was higher in shade than in sun, and in high than in low irrigation, while Q. faginea – from more humid zones – responded to irrigation only. Shade-induced growth was accompanied by a decline in N concentration in the evergreens, but irrigation reduced N concentration only of Q. faginea. Shade, but not irrigation, reduced above-ground N loss. We conclude that both treatments differentially affected the evergreen and the deciduous oaks, probably due to differences in plant hydraulic and stomatal conductance. Although both treatments have similar effects on the growth of evergreens, they produced different effects on seedling N economy, which may have important consequences on future field seedling performance.     

Effectiveness of rodents as local seed dispersers of Holm oaks

In this study we assessed the effectiveness of rodents as dispersers of Quercus ilex in a patchy landscape in southeastern Spain. We experimentally followed the fates of 3,200 marked and weighed acorns from dispersal through the time of seedling emergence over three years. Rodents handled about 99% of acorns, and dispersed 67% and cached 7.4% of the dispersed acorns. Most caches were recovered and consumed, and only 1.3% of the original experimental acorns were found alive in caches the following spring. Dispersal distances were short (mean = 356.2 cm, median = 157 cm) and strongly right-skewed. Heavier acorns were dispersed further and were more likely to be cached and survive than lighter acorns. All caches were in litter or soil, and each contained a single acorn. Rodents moved acorns nonrandomly, mostly to oaks and pines. Most surviving acorns were either in oaks, a poor microhabitat for oak recruitment, or shrubs, a suitable microhabitat for oak recruitment. Our results suggest that rodents, by burying a relatively high proportion of acorns singly in shrubs and pines, act as moderately effective dispersers of Q. ilex. Nonetheless, this dispersal comes at a very heavy cost.     

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.     

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.     

Hoarding behaviors of two wood mouse species: Different preference for acorns of two Fagaceae species

Two species of wood mouse, Apodemus argenteus and A. speciosus, were observed consuming and hoarding acorns of Quercus serrata and Castanopsis cuspidata. When each species of acorn was supplied individually, both species of mice used each species of acorn for eating and hoarding. When both species of acorn were supplied, A. argenteus consumed or hoarded only C. cuspidata, whereas A. speciosus tended to eat C. cuspidata acorns at the feeding site, and disperse or hoard Q. serrata acorns. Apodemus speciosus is unlikely to disperse C. cuspidata acorns (their utilization was biased towards consumption) when Q. serrata acorns are also available. Apodemus argenteus will make almost no contribution to the dispersal of Q. serrata when the two acorn species coexist.     

Guild structures of seed insects in relation to acorn development in two oak species

We investigated patterns of acorn growth in Quercus variabilis Blume and Quercus serrata Thunb., seasonal trends in emergence of seed insects found in the acorns, oviposition periods of the insects and falling periods of insect-infested acorns. In Q. variabilis, two insect guilds were associated with acorn development: (i) the immature acorn feeding (IAF) guild [Curculionidae sp., Poecilips cardamomi (Schaufuss), and Characoma ruficirra (Hampson)]; and (ii) the mature acorn feeding (MAF) guild [Curculio robustus (Roelofs), Curculio sikkimensis (Heller), and Cydia glandicolana (Danilevsky)]. In Q. serrata, there were three guilds: (i) the pistillate flower feeding (PFF) guild (cynipid wasp); (ii) the IAF guild [sap absorption by Mechoris ursulus (Roelofs)]; and (iii) the MAF guild (M. ursulus, C. sikkimensis, Cydia danilevskyi (Kuznetzov), C. glandicolana and Autostichidae sp.). The succession of guilds during acorn development may be a consequence of the use by different species of the limited food resource. The lack of a PFF guild in Q. variabilis that was found in our field site would have a positive effect on IAF guilds in utilizing the acorns.     

Root Morphology, Stem Growth and Field Performance of Seedlings of Two Mediterranean Evergreen Oak Species Raised in Different Container Types

Outplanting container-grown oak seedlings with undesirable shoot and root characteristics result in poor establishment and reduced field growth. The objective of this study was to determine the influence of container type on both above-and below-ground nursery growth and field performance of one-year old tap-rooted seedlings Quercus ilex L. and Quercus coccifera L. The experiment was conducted in an open-air nursery and the seedlings were grown in three container types. At the end of the nursery, growth period seedlings’ shoot height, diameter (5 mm above root collar), shoot and root biomass, root surface area, root volume and total root length were assessed. Then the seedlings were planted in the field and their survival and growth were recorded for two growing seasons after outplanting. The results showed a difference between the Quercus species in the effect of container type. Q. ilex seedlings raised in paper-pot had significantly greater height, diameter, shoot and root biomass and root volume than those raised in the other two container types. Similarly, Q. coccifera seedlings raised in paper-pot, had significantly greater above-and below-ground growth than those raised in the other two container types. Both oak species showed relatively low survival in the field; the mortality was mainly observed the first year after outplanting, especially after the summer dry period. However, 2 years after outplanting, the paper-pot seedlings of the two oak species showed better field performance.     

Effects of acorn masting on population dynamics of three forest-dwelling rodent species in Hokkaido,Japan

The effects of the abundance of acorns of the oak, Quercus crispula, on the population dynamics of three rodent species (Apodemus speciosus, A. argenteus, and Clethrionomys rufocanus) were analyzed using time series data (1992–2006). The data were obtained in a forest in northern Hokkaido, Japan, by live trapping rodents and directly counting acorns on the ground. Apodemus speciosus generally increased in abundance following acorn masting. However, the clear effect of acorn abundance was not detected for the other two rodent species. Acorns of Q. crispula contain tannins, which potentially have detrimental effects on herbivores. Apodemus speciosus may reduce the damage caused by acorn tannins with tannin-binding salivary proteins and tannase-producing bacteria, whereas such physiological tolerance to tannins is not known in the other two rodent species. The differences in the effects of acorns between the three species may be due to differences in their physiological tolerance to tannins.     

Acorn crop size and pre-dispersal predation determine inter-specific differences in the recruitment of co-occurring oaks

The contribution of pre-dispersal seed predation to inter-specific differences in recruitment remains elusive. In species with no resistance mechanisms, differences in pre-dispersal predation may arise from differences in seed abundance (plant satiation) or in the ability of seeds to survive insect infestation (seed satiation). This study aimed to analyse the impact of pre-dispersal acorn predation by weevils in two co-occurring Mediterranean oaks (Quercus ilex and Quercus humilis) and to compare its relevance with other processes involved in recruitment. We monitored the patterns of acorn production and acorn infestation by weevils and we conducted experimental tests of acorn germination after weevil infestation, post-dispersal predation and seedling establishment in mixed forests. Monitoring and experimental data were integrated in a simulation model to test for the effects of pre-dispersal predation in recruitment. In both oaks pre-dispersal acorn infestation decreased with increasing acorn crop size (plant satiation). This benefited Q. ilex which exhibited stronger masting behaviour than Q. humilis, with almost a single and outstanding reproductive event in 6 years. Acorn infestation was more than twice as high in Q. humilis (47.0%) as in Q. ilex (20.0%) irrespective of the number of seeds produced by each species. Although germination of infested acorns (seed satiation) was higher in Q. humilis (60%) than in Q. ilex (21%), this could barely mitigate the higher infestation rate in the former species, to reduce seed loss. Conversely to pre-dispersal predation, no inter-specific differences were observed either in post-dispersal predation or seedling establishment. Our results indicate that pre-dispersal predation may contribute to differences in seed supply, and ultimately in recruitment, between co-existing oaks. Moreover, they suggest that seed satiation can barely offset differences in seed infestation rates. This serves as a warning against overemphasising seed satiation as a mechanism to overcome seed predation by insects. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Dietary circumvention of acorn tannins by blue jays

Blue jays consume large quantities of acorns to fuel energy-demanding caching flights in the fall. Yet blue jays possess no known physiological adaptation to counter the negative effects of a high tannin diet on protein digestion. Dietary experiments were conducted to determine if blue jays could subsist on an acorn-only diet, and if they could not, to determine whether supplements of acorn weevil larvae (Curculio), present inside acorns, enabled them to maintain their mass. Comparative tannin assays also were conducted on Lepidobalanus (low tannin; white oak) and Erythrobalanus (high tannin; pin oak) acorns using radial diffusion assay. Captive jays consumed considerable acorn material, yet were unable to maintain mass on ad lib. acorn-only diets or on an acorn +1.5 g larvae/day supplement. There were no significant differences in mass loss between high and low tannin diets. In contrast, blue jays were able to stabilize mass on a diet of acorns +5.0 g larvae supplement/day. These results suggest that acorn weevil larvae, or perhaps other insects, counteract the effects of acorn tannins in the jay diet allowing jays to subsist largely on acorns during the fall caching season. Oak demographic processes may be partly regulated by a tri-trophic relationship among plant, insect and bird. Acorn weevil larvae, considered damaging to oak populations, may actually facilitate oak recruitment and population vagility in the long-term.     

Cytosolic ascorbate peroxidase in some species of the Quercus genus

ABSTRACT

Six species of the Quercus genus (Quercus ilex L., Q. coccifera L., Q. suber L., Q. trojana Webb, Q. macrolepis Kotschy, Q. cerris L.) have been screened for cytosolic ascorbate peroxidase (APX) by means of native polyacrylamide gel electrophoresis (PAGE). A single isozyme was found in five species (Q. trojana, Q. suber, Q. cerris, Q. macrolepis and Q. coccifera), while Q. ilex shows two different APX proteins. The data showed marked similarities among Q. trojana, Q. suber, Q. cerris and Q. macrolepis with respect to the electrophoretic mobility. The validity of APX electrophoretic patterns in systematic studies is discussed.     

Acorn production is linked to secondary growth but not to declining carbohydrate concentrations in current-year shoots of two oak species

In trees, reproduction constitutes an important resource investment which may compete with growth for resources. However, detailed analyses on how growth and fruit production interact at the shoot level are scarce. Primary canopy growth depends on the development of current-year shoots and their secondary growth might also influence the number and size of fruits supported by them. We hypothesise that an enhanced thickening of current-year shoots is linked positively to acorn production in oaks. We analysed the effect of acorn production on shoot growth of two co-occurring Mediterranean oak species with contrasting leaf habit (Quercus ilex, Quercus faginea). Length and cross-sectional area of current-year shoots, apical bud mass, number of leaves and acorns, xylem and conductive area, number of vessels of acorn-bearing and non-bearing shoots were measured in summer and autumn. Nitrogen and carbohydrates analyses were also performed in stems and leaves of both shoot types. Stem cross-sectional area increased in acorn-bearing shoots when compared with non-bearing shoots for both species and such surplus secondary growth was observed since summer. In bearing shoots, the total transversal area occupied by vessels decreased significantly from basal to apical positions along the stem as did the xylem area and the number of vessels. Leaves of bearing shoots showed lower nitrogen concentration than those of non-bearing shoots. Carbohydrate concentrations did not differ in stems and leaves as a function of the presence of acorns. Such results suggest that carbohydrates may preferentially be allocated towards reproductive shoots, possibly through enhanced secondary growth, satisfying all their carbon demands for growth and reproduction. Our findings indicate that acorn production in the two studied oaks depends on shoot secondary growth.     

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.     

Quercus ilex facing water stress: a functional equilibrium hypothesis

A boundary line analysis of the scatterplot relating pre-dawn leaf water potential to pre-dawn minus minimum leaf water potential was applied to study the efficiency of the Mediterranean evergreen oak species to buffer the effects of variability in water resources. The results are discussed in term of stomatal regulation and changes in soil-to-leaf hydraulic conductance of water flow, both induced by changes in leaf water potential. For this purpose, we used data for leaf water potential measured in three stands of Quercus ilex across a soil water availability gradient in Southern France, and two others obtained in California and Arizona for Quercus turbinella and Portugal for Quercus coccifera. A classification of plant responses from mesic to xeric sites is proposed.