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.     

Litter dynamics in post-fire successional forests of Quercus ilex

Forest floor regeneration after fire in a holm oak forest was studied in a chronosequence in Catalonia (NE Spain). The sampled area represented situations of high fire disturbance because of the frequency of wildfires and the high impact of erosive processes. Forest floor bulk samples from the L, F and H layers were taken in 5 burned plots, aging from 0 to 35 years. They were analyzed for dry standing weight, and organic carbon and N content.Plant cover reached almost 100% two years after the fire. During the first 20 years, shrubs and herbs had the highest surface covering, and thereafter, holm oaks became dominant.Holm oak litterfall stabilized by the fourth year after burning.The major shrub species which were present in the chronosequence had two different patterns with respect to nitrogen: 1) a low litter C/N ratio (i.e. nitrogen fixing species); 2) a high litter C/N ratio.The litter decay coefficients were relatively high, allowing for a rapid structuration and formation of the L and F layers after the fire. Indeed, 95% of the maximum steady standing weight accumulates in 8–9 years. The pattern of cover in the different strata influenced the lateral distribution and variable accumulation of the forest floor layers and determined different turnover rates.     

Effect of water stress on monoterpene emissions from young potted holm oak (Quercus ilex L.) trees

We investigated the effects of a short period of water stress on monoterpene emissions from Quercus ilex, a common oak species of the Mediterranean vegetation and a strong emitter of monoterpenes. The experiment was carried out on two young saplings with a branch enclosure system under semi-controlled conditions. Under unstressed conditions, small qualitative (cis--ocimene, trans--ocimene, -caryophyllene and 1,8-cineol) and large quantitative (as much as 40% for the main compounds emitted) differences were observed between the two apparently similar trees. Nevertheless these differences did not affect the short-and long-term responses to temperature and water stress. Daily courses of emissions and gas exchanges were similar before and after the stress. During the most severe stress, emissions were reduced by a factor of two orders of magnitude and the log-linear relationship between emissions and temperature no longer existed. Photosynthesis and transpiration rates decreased as soon as the soil started to dry, whereas monoterpene emissions slightly increased for few days and then dropped when the daily CO₂ balance approached zero. We concluded that under water stress monoterpene emissions were highly limited by monoterpene synthesis resulting from a lack of carbon substrate and/or ATP. After rewatering, both emissions and gas exchanges recovered immediately, but to a level lower than the pre-stress level. These results have many implications for monoterpene emission modelling in the Mediterranean area, since the dry period generally extends from May to August. If our results are confirmed by field experiments, water stress could lead to a large overestimation of the emissions under summer conditions, when the algorithms based on light and temperature would give high emission rates.     

Survival and growth of konara oak (Quercus serrata Thunb.) seedlings in an abandoned coppice forest

Survival and growth of konara oak (Quercus serrata) seedlings were examined on the forest floor of a konara oak coppice stand in Mitaka, Tokyo, for 5 yr (1976–1980) after exclusion of human management. Attention was mainly focused on a large cohort derived from a bumper acorn crop in 1975. Seedlings began to branch in the second year, although some did not branch during the whole 5-yr period. Branched seedling showed higher survival and greater weight in 1980 than unbranched seedlings. Many of the branched seedlings had lost an orthotropic leader shoot and spread their branches laterally, suggesting a response for efficient light-capture. As for unbranched seedlings, a size relationship between leaf area and stem length developing every year was dependent on the shoot size. A small shoot gave priority to enlargement of leaf area, whereas a large shoot gave priority to stem elongation. Yearly variations in seedling mortality and stem elongation were related to the amount of precipitation. Both the initial height and leaf area of germinated seedlings were positively correlated with their survival and weight after five years. Five-year survival of seedlings showed a positive correlation with light intensity at the growth site but a negative correlation with the extent of leaf predation. Sprouts of shrub species grew vigorously and became much taller than the oak seedlings in the 5-yr observation period. The characteristics of forest-floor seedlings were discussed in relation to the regeneration of konara oak forests in general.     

Leaf flavonoid content in Quercus ilex L. resprouts and its seasonal variation

Here, we provide the first report on flavonoid content in holm oak (Quercus ilex L.) leaves, analyzed by HPLC–MS/MS. Flavanols and flavonols were the predominant groups, although proanthocyanidins and many soluble tannins had a relevant presence in all leaf samples. Seasonal variation of flavonoids was determined in extracts from Q. ilex leaves during resprouting after a forest fire in two Mediterranean forests. Similar seasonal trends were observed over 2 years during the two main stress seasons (winter and summer). The most abundant flavonoid was the flavanol epicatechin, which showed similar values during the two seasons. Hexosides of the flavonols, quercetin, kaempferol and rhamnetin showed considerably higher content in winter, especially at the lowest temperatures. These variations in both forests are discussed on the basis of the chlorophyll fluorescence results obtained. Anthocyanins were found practically absent in mature leaves. Nutrient or water availability differences between sites or seasons were not related to changes in leaf flavonol-hexoside content.     

The facultative bimodal growth pattern in Quercus ilex – A simple model to predict sub-seasonal and inter-annual growth

In Mediterranean climates, bimodal growth patterns, corresponding to two peaks in radial increment during favorable seasons, have been described in several tree species. However, we lack a better mechanistic understanding of bimodality and its potential responses to the predicted warming and aridification trends. Filling this research gap is important since growth duration affects the capacity of trees to form wood and uptake carbon. Here we used an 11-year (1994–2004) long record of dendrometer data of the Mediterranean Holm oak (Quercus ilex) and compared how climate related to radial increment in trees from the south- and the north-facing slopes. We also related climate variables to tree-ring width and the production of intra-annual density fluctuations (IADFs), which reflects bimodality. In this paper, we introduce a model called VS-Lite2 to simulate tree-growth dynamics, which is a modified version of the process-based Vaganov-Shashkin Lite model. The VS-Lite2 model adequately reproduced the bimodal intra‐annual pattern of radial growth, IADFs, and annual tree growth. Trees from the south-oriented slope grew more, produced more IADFs and showed a more marked bimodal pattern than trees from the north-facing slope. These differences agree with the observation that late-summer drought constrained growth. Therefore, radial-growth models should consider plastic bimodality and micro-environmental conditions in areas subjected to seasonal droughts.     

Germination of Quercus ilex and Q. pubescens in a Q. ilex coppice

Quercus ilex L. (holm oak) coppices, widespread around the Mediterranean basin, are probably the result of 5 000 years of prolonged human disturbance of the original Quercus pubescens Willd. (downy oak) forests. Since disturbance has almost ceased in recent years, a question arises as to the development of these coppices: will the Q. pubescens forests return, or will Q. ilex remain the dominant species? To investigate the phenomenon, we analyzed the first stages, i.e. germination of the two species in holm oak coppices. Our experiments show that both species germinated better in coppices than in clearings or clear-cuts. Moreover, Q. pubescens appears to be slightly favored above Q. ilex and it is suggested auto-allelopathy is involved, at least partially inhibiting the germination of Q. ilex.     

Light and temperature dependence of the emission of cyclic and acyclic monoterpenes from holm oak (Quercus ilex L.) leaves

In a laboratory study, we investigated the monoterpene emissions from Quercus ilex, an evergreen sclerophyllous Mediterranean oak species whose emissions are light dependent. We examined the light and temperature responses of individual monoterpenes emitted from leaves under various conditions, the effect of heat stress on emissions, and the emission-onset during leaf development. Emission rate increased 10-fold during leaf growth, with slight changes in the composition. At 30 °C and saturating light, the monoterpene emission rate from mature leaves averaged 4·1 nmol m^–2 s^–1, of which α-pinene, sabinene and β-pinene accounted for 85%. The light dependence of emission was similar for all monoterpenes: it resembled the light saturation curve of CO₂ assimilation, although monoterpene emission continued in the dark. Temperature dependence differed among emitted compounds: most of them exhibited an exponential increase up to 35 °C, a maximum at 42 °C, and a slight decline at higher temperatures. However, the two acyclic isomers cis-β-ocimene and trans-β-ocimene were hardly detected below 35 °C, but their emission rates increased above this temperature as the emission rates of other compounds fell, so that total emission of monoterpenes exponentially increased from 5 to 45 °C. The ratio between ocimene isomers and other compounds increased with both absolute temperature and time of heat exposure. The light dependence of emission was insensitive to the temperature at which it was measured, and vice versa the temperature dependence was insensitive to the light regime. The results demonstrated that none of the models currently applied to simulate isoprene or monoterpene emissions correctly predicts the short-term effects of light and temperature on Q. ilex emissions. The percentage of fixed carbon lost immediately as monoterpenes ranged between 0·1 and 6·0% depending on temperature, but rose up to 20% when leaves were continuously exposed to temperatures between 40 and 45 °C.     

The effect of low temperatures on the photosynthetic apparatus of Quercus ilex subsp. ballota at its lower and upper altitudinal limits in the Iberian peninsula and during a single freezing-thawing cycle

We investigated the response of the photosynthetic apparatus during an episode of extreme low winter temperature in Quercus ilex subsp. ballota (Desf.) Samp., a typical Mediterranean evergreen species in the Iberian peninsula. Both plants in a woodland located at high altitude (1,177 m. a.s.l.) and potted plants obtained from acorns of the same populations grown at low altitude (225 m. a.s.l.) were analyzed. Net CO₂ assimilation rate was negative and there was a marked decrease in photosystem II (PSII) efficiency during winter in leaves of the woodland population (high altitude individuals). These processes were accompanied by increases in non-photochemical quenching (NPQ) and in the de-epoxidated carotenoids within the xanthophyll cycle, mechanisms aimed to dissipate excess energy. In addition, these de-epoxidated carotenoids were largely preserved during the night. There was no chlorophyll bleaching during the winter, which suggests that leaves were not experiencing photoinhibitory damage. In fact, the net photosynthetic rate and the PSII efficiency recovered in spring. These changes were not observed, or were much more reduced, in individuals located at lower altitude after a few frosts. When the response to rapid temperature changes (from 20°C to –5°C and from –5°C to 20°C) was studied, it was found that the maximum potential PSII efficiency was fairly stable, ranging from 0.70 to 0.75. The rest of the photosynthetic parameters measured, actual and intrinsic PSII efficiency, photochemical and NPQ, responded immediately to the changes in temperature and, also, the recovery after cold events was practically immediate.     

Drought stress does not protect Quercus ilex L. from ozone effects: results from a comparative study of two subspecies differing in ozone sensitivity

Long‐term effects of ozone (O₃) exposure and drought stress were assessed on two subspecies of Quercus ilex: ssp. ilex and ssp. ballota. Two‐year‐old seedlings were continuously exposed for 26 months in open‐top chambers to three O₃ treatments: charcoal filtered air, non‐filtered air and non‐filtered air supplemented with 40 nl·l^?1 O₃. Additionally, two irrigation regimes were adopted: half of the plants were well‐watered and the others received half of the water supplied to control plants. Growth, shoot water potential and gas exchange rates were assessed seasonally, and biomass accumulation was determined at the end of the experiment. Drought stress caused higher reductions of gas exchange, growth and biomass accumulation than O₃ exposure in both subspecies. The combination of O₃ and drought stress caused further decreases of accumulated aboveground biomass but no additive effects were observed on gas exchange rates or root biomass. Thus, drought stress did not protect Q. ilex from O₃ effects on biomass when the response of the whole plant was considered. Q. ilex ssp. ballota was more sensitive to O₃ and ssp. ilex was more affected by drought stress. The different O₃ sensitivity was not only related to pollutant uptake but also to the ability of plants for resource acquisition and allocation. Based on biomass dose–response functions, Q. ilex is more resistant to O₃ than other European evergreen tree species, however, O₃ represents an additional stress factor that might be impairing plant ability to withstand current and future climate change.     

Regeneration by sprouting of holm-oak (Quercus ilex) stands exploited by selection thinning

Holm-oak forests of the Montseny Massif have traditionally been exploited by selection thinning, which consists of a partial removal of standing trees and biomass. Cutting percentages within each size class indicate that this management technique represents a compromise between forest improvement and harvesting.Sprouting is the mean regeneration process of these holm-oak stands. The mean number of living sprouts per stool is considerably higher than in other coppiced hardwood species, and decreases along the cutting-cycle according to a negative exponential function. Mean number of surviving sprouts per stool is significantly correlated with regenerative age and stool size. Death of sprouts throughout the cutting cycle is compensated by growth of surviving sprouts. Thus, the weight of living sprouts per stool increases linearly with time, and also with the basal area coppiced.Sprouts developing within the first fifteen years of regeneration belong to a single cohort that appears during the first year after cutting. After that period of time, new sprouts appear from dormant buds, so that after 30 years of regeneration a wide range of cohorts of different ages are present.     

Annual and seasonal changes in fine root biomass of a Quercus ilex L. forest

The biomass, production and mortality of fine roots (roots with diameter <2.5 mm) were studied in a typical Mediterranean holm oak (Quercus ilex L.) forest in NE Spain using the minirhizotron methodology. A total of 1212 roots were monitored between June of 1994 and March of 1997. Mean annual fine root biomass in the holm oak forest of Prades was 71±8 g m^–2 yr^–1. Mean annual production for the period analysed was 260+11 g m^–2 yr^–1. Mortality was similar to production, with a mean value of 253±3 g m^–2 yr^–1. Seasonal fine root biomass presented a cyclic behaviour, with higher values in autumn and winter and lower in spring and summer. Production was highest in winter, and mortality in spring. In summer, production and mortality values were the lowest for the year. Production values in autumn and spring were very similar. The vertical distribution of fine root biomass decreased with increasing depth except for the top 10–20 cm, where values were lower than immediately below. Production and mortality values were similar between 10 and 50 cm depth. In the 0–10 cm and the 50–60 cm depth intervals, both production and mortality were lower.     

Morphological adaptions of Quercus ilex leaves in the Castelporziano forest

The annual course of vegetative growth of the most representative species of a Quercus ilex L. forest in Castelporziano (Latium, Italy) was studied through periodical analysis of selected ecophysiological leaf indexes, for the period 1987–1990. The results demonstrate that the local climate facilitates continuous vegetative activity of the sclerophyllous species without a latent phase. The variable morphology of sun and shade leaves of Q. ilex illustrates the plant's response to environmental stress.     

Carbon isotope discrimination and wood anatomy variations in mixed stands of Quercus robur and Quercus petraea

The two most common oak species in western Europe, Quercus robur and Quercus petraea, display different ecological behaviours, particularly with respect to their responses to drought. The ecophysiological basis of this niche difference is not understood well. Here we test the hypothesis that these two species present distinct water use efficiencies (WUEs), using the carbon isotope discrimination approach. Leaves and 13 dated ring sequences were sampled in 10 pairs of adult trees growing side by side. Carbon isotope composition was measured on cellulose extracts. In addition, relationships between carbon isotope discrimination and wood anatomy were assessed at the tree level. Quercus robur displayed a 1·0‰ larger isotopic discrimination than Q. petraea, and therefore a lower intrinsic WUE (?13%). This interspecific difference of isotopic discrimination was quite stable with time and independent of tree radial growth and climate fluctuations. A strong positive correlation was observed between average tree values of earlywood vessel surface area and ¹³C isotopic discrimination. This correlation was even higher with ¹³C of the 1976 dry year (r = 0·86). These observations led to the hypothesis that hydraulic properties of xylem could exert a constraint on leaf gas exchange, resulting in a larger WUE for individuals with smaller vessel cross‐section area.     

Isoprenoid emission in trees of Quercus pubescens and Quercus ilex with lifetime exposure to naturally high CO2 environment†

The long‐term effect of elevated atmospheric CO₂ on isoprenoid emissions from adult trees of two Mediterranean oak species (the monoterpene‐emitting Quercus ilex L. and the isoprene‐emitting Quercus pubescens Willd.) native to a high‐CO₂ environment was investigated. During two consecutive years, isoprenoid emission was monitored both at branch level, measuring the actual emissions under natural conditions, and at leaf level, measuring the basal emissions under the standard conditions of 30 °C and at light intensity of 1000 µmol m^?2 s^?1. Long‐term exposure to high atmospheric levels of CO₂ did not significantly affect the actual isoprenoid emissions. However, when leaves of plants grown in the control site were exposed for a short period to an elevated CO₂ level by rapidly switching the CO₂ concentration in the gas‐exchange cuvette, both isoprene and monoterpene basal emissions were clearly inhibited. These results generally confirm the inhibitory effect of elevated CO₂ on isoprenoid emission. The absence of a CO₂ effect on actual emissions might indicate higher leaf temperature at elevated CO₂, or an interaction with multiple stresses some of which (e.g. recurrent droughts) may compensate for the CO₂ effect in Mediterranean ecosystems. Under elevated CO₂, isoprene emission by Q. pubescens was also uncoupled from the previous day's air temperature. In addition, pronounced daily and seasonal variations of basal emission were observed under elevated CO₂ underlining that correction factors may be necessary to improve the realistic estimation of isoprene emissions with empirical algorithms in the future. A positive linear correlation of isoprenoid emission with the photosynthetic electron transport and in particular with its calculated fraction used for isoprenoid synthesis was found. The slope of this relationship was different for isoprene and monoterpenes, but did not change when plants were grown in either ambient or elevated CO₂. This suggests that physiological algorithms may usefully predict isoprenoid emission also under rising CO₂ levels.     

Diversity of saproxylic basidiomycetes in Quercus ilex woodlands of central and insular Greece

The sclerophyllous forests of Quercus ilex (holm oak) possess a unique position in earth’s biosphere, existing only in countries of the Mediterranean basin. However, very little information exists about mushrooms associated with holm oak in east Mediterranean, including Greece, where Q. ilex appears mostly in relict forests and fragmented stands. A pertinent long-term investigation was undertaken in selected Q. ilex dominated habitats of Crete, Andros, Naxos and Ikaria islands (Aegean Archipelago) as well as in mountains of Attica. Specimens of xylotrophic basidiomycetes were collected and identified. As a result, 74 species were recorded; among them Hemimycena cephalotricha, Hyphoderma cremeoalbum, Hyphodontia radula, Irpex litschaueri, Mycena algeriensis, Phanerochaete martelliana, Phloeomana alba and Phlebia unica constitute new records for the Greek mycobiota, while 68 species are reported for the first time on Q. ilex in Greece. Moreover, the presence of Fomitiporia mediterranea on this host was evidenced through ITS sequencing, and comments are made about its relationship to F. punctata and F. pseudopunctata. The importance of findings is discussed in conjunction with their environmental value; five species could be assigned in the “Species of Special Interest” categories, and other five polypores are proposed as suitable indicators for Q. ilex habitat assessment and conservation.     

Comparison of earlywood vessel variables in the wood of Quercus robur L. and Quercus petraea (Mattuschka) Liebl. growing at the same site

The differences in the microscopic structure of wood based on the variables of earlywood vessel area and tree ring width were analysed in 6 trees of Pedunculate oak (Quercus robur L.) and 6 trees of Sessile oak (Quercus petraea (Mattuschka) Liebl.) in the same forest stand at a site in the Vizovice Highland (Czech Republic). The aim of this paper was to assess any differences between the two oak species when grown in the same location. Also the effect of tree-ring width and the effect of age on earlywood vessel variables were analysed. Mean values of tree-ring widths were not different between species. Earlywood vessel area chronologies were synchronized well between species. The mean values of earlywood vessel area (average vessel area, average vessel area of the first row of vessels, area of the largest vessel) showed a significant difference. Finally, we can conclude that there were differences in vessel features between these oak species.     

The role of disturbance in the co‐existence of the evergreen Quercus ilex and the deciduous Quercus cerrioides

Question: Which is the response of the evergreen Quercus ilex and the deciduous Q. cerrioides to repeated disturbances? Location: central Catalonia (northeastern Spain), in the areas affected by two of the largest historically recorded wildfires in NE Spain: the Bages‐Berguedà fire (24 300 ha forested area burned in July 1994), and the Solsonès fire (14 300 ha burned in 1998). Methods: Survival and growth of individuals of Quercus ilex and Q. cerrioides were evaluated in plants subjected to different fire histories and experimental disturbances (burning, cutting or clipping) applied either before or after summer. Results: Survival was high (> 99%), with both species showing a similar high resistance to disturbances. Growth after experimental disturbance was positively related to the size of the individual before the latest forest fire occurred. Fire history had a large effect on resprout growth, as the repeated incidence of disturbances lowered the capacity of individuals to grow. The type and season of experimental disturbance experienced by plants had a large effect. Individuals that experienced total above‐ground loss had lower growth rates than those with partial loss. A similar pattern was observed in individuals disturbed after the summer in relation to those disturbed before summer. Conclusions: The larger growth rates recorded in Q. cerrioides across all fire histories and experimental treatments, and the higher vulnerability of Q. ilex to increased fire frequency, intensity of experimental disturbance, and disturbance season, provide evidence for the relatively high susceptibility of the latter to repeated disturbances. This view disagrees with the larger resilience of this species compared to co‐existing deciduous oaks, as reported.