Topography modulates climate sensitivity of multidecadal trends of holm oak decline

Forest decline events have increased worldwide over the last decades being holm oak (Quercus ilex L.) one of the tree species with the most worrying trends across Europe. Since this is one of the tree species with the southernmost distribution within the European continent, its vulnerability to climate change is a phenomenon of enormous ecological importance. Previous research identified drought and soil pathogens as the main causes behind holm oak decline. However, despite tree health loss is a multifactorial phenomenon where abiotic and biotic factors interact in time and space, there are some abiotic factors whose influence has been commonly overlooked. Here, we evaluate how land use (forests versus savannas), topography, and climate extremes jointly determine the spatiotemporal patterns of holm oak defoliation trends over almost three decades (1987–2014) in Spain, where holm oak represents the 25% of the national forested area. We found an increasing defoliation trend in 119 out of the total 134 holm oak plots evaluated, being this defoliation trend significantly higher in forests compared with savannas. Moreover, we have detected that the interaction between topography (which covariates with the land use) and summer precipitation anomalies explains trends of holm oak decline across the Mediterranean region. While a higher occurrence of dry summers increases defoliation trends in steeper terrains where forests dominate, an inverse relationship was found in flatter terrains where savannas are mainly located. These opposite relationships suggest different causal mechanisms behind decline. Whereas hydric stress is likely to occur in steeper terrains where soil water holding capacity is limited, soil waterlogging usually occurs in flatter terrains what increases tree vulnerability to soil pathogens. Our results contribute to the growing evidence of the influence of local topography on forest resilience and could assist in the identification of potential tree decline hotspots and its main causes over the Mediterranean region.     

Landscape dynamics in Mediterranean oak forests under global change: understanding the role of anthropogenic and environmental drivers across forest types

The Mediterranean region is projected to be extremely vulnerable to global change, which will affect the distribution of typical forest types such as native oak forests. However, our understanding of Mediterranean oak forest responses to future conditions is still very limited by the lack of knowledge on oak forest dynamics and species‐specific responses to multiple drivers. We compared the long‐term (1966–2006) forest persistence and land cover change among evergreen (cork oak and holm oak) and deciduous oak forests and evaluated the importance of anthropogenic and environmental drivers on observed changes for Portugal. We used National Forest Inventories to quantify the changes in oak forests and explored the drivers of change using multinomial logistic regression analysis and an information theoretical approach. We found distinct trends among oak forest types, reflecting the differences in oak economic value, protection status and management schemes: cork oak forests were the most persistent (62%), changing mostly to pines and eucalypt; holm oak forests were less persistent (53.2%), changing mostly to agriculture; and deciduous oak forests were the least persistent (45.7%), changing mostly to shrublands. Drivers of change had distinct importance across oak forest types, but drivers from anthropogenic origin (wildfires, population density, and land accessibility) were always among the most important. Climatic extremes were also important predictors of oak forest changes, namely extreme temperatures for evergreen oak forests and deficit of precipitation for deciduous oak forests. Our results indicate that under increasing human pressure and forecasted climate change, evergreen oak forests will continue declining and deciduous oak forests will be replaced by forests dominated by more xeric species. In the long run, multiple disturbances may change competitive dominance from oak forests to pyrophytic shrublands. A better understanding of forest dynamics and the inclusion of anthropogenic drivers on models of vegetation change will improve predicting the future of Mediterranean oak forests.     

Tree growth,mortality, and above-ground biomass accumulation in a holm oak forest under a five-year experimental field drought

A holm oak forest was exposed to an experimental drought during 5 years to elucidate the growth responses of the dominant species Quercus ilex, Arbutus unedo and Phillyrea latifolia. Soil water availability was partially reduced, about 15% as predicted for this area for the next decades by GCM and ecophysiological models, by plastic strips intercepting rainfall and by ditch exclusion of water runoff. The stem diameter increment was highly correlated with annual rainfall in all species, and drought treatment strongly reduced the diameter increment of Q. ilex (41%) and specially of A. unedo (63%), the species showing higher growth rates. Stem mortality rates were highly correlated with previous stem density, but drought treatment increased mortality rates in all species. Q. ilex showed the highest mortality rates (9% and 18% in control and drought plots, respectively), and P. latifolia experienced the lowest mortality rates (1% and 3% in control and drought plots, respectively). Drought strongly reduced the increment of live aboveground biomass during these 5 years (83%). A. unedo and Q. ilex experienced a high reduction in biomass increment by drought, whereas P. latifolia biomass increment was insensitive to drought. The different sensitivity to drought of the dominant species of the holm oak forest may be very important determining their future development and distribution in a drier environment as expected in Mediterranean areas for the next decades. These drier conditions could thus have strong effects on structure (species composition) and functioning (carbon uptake and biomass accumulation) of these Mediterranean forests.     

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

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

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

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

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.     

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.     

Modelling Mediterranean oak palaeolandscapes using the MaxEnt model algorithm: The case of the NE Iberia under the Middle Holocene climatic scenario

The Mediterranean Basin is a global biodiversity hotspot, and oak tree species play an important role in it. Since the beginning of the Holocene (∼11.4 kyr BP), the distribution of forests has not occurred uniformly, resulting in diverse types of vegetation landscapes. In this study, we used a maximum entropy algorithm (MaxEnt) to obtain the ecological niche model (ENM) of two sub-Mediterranean oak species, Quercus pubescens Willd. (pubescent oak) and Quercus ilex subsp. ilex (holm oak), both in the present day in the Iberian Peninsula and within a Middle Holocene (8.2–4.2 kyr BP) climatic scenario in the NE Iberian Peninsula. Moreover, we used the locations of Neolithic archaeological sites containing anthracological data to analyze the relationship between human occupations and oak habitats. Our results suggest that the two oaks have responded differently to the climatic conditions that have occurred, and show changes in both potential distributions. The palaeolandscape vegetation map shows a denser vegetation cover in the lowlands and a more open landscape in the highlands, with a higher dominance of Quercus pubescens in the septentrional areas, while Quercus ilex was more restricted to certain coastal areas. Temperature and precipitation factors, mainly seasonal climatic conditions, have had a greater impact on the distribution of vegetation than other factors. We found a good overlap between the ENM of the two oaks and the locations of the Neolithic sites analysed, and determined that the distribution of Neolithic archaeological sites is not random. The Neolithic populations in the study area depended heavily on the resources of the deciduous and evergreen sub- Mediterranean forest, although they also exploited the resources of the mountain pine forest. Neolithic sites distribution suggests that Neolithic human groups were aware of the potential of forests and probably gathered woody resources in their surroundings.     

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.     

Evaluating the benefits of CAP reforms: Can afforestations restore bird diversity in Mediterranean Spain?

Current Common Agricultural Policy (CAP) subsidies in the Mediterranean region tend to prioritize afforestation on former arable land with oaks rather than pines because pine plantations would maintain lower biological diversities than native forests. Nevertheless, no thorough evaluations of the conservation values of pine plantations as compared to oak remnants have been carried out to date. We analyze the diversity and conservation value of bird assemblages breeding in 200 remnants of Holm oak Quercus ilex woodlands and 82 mature (>50-year-old) pine plantations in central Spain, a Mediterranean region mostly devoted to arable farming. Species–area relationships were compared between forest types. The conservation value of bird assemblages was assessed using the “Species of European Conservation Concern” (SPEC) classification of Burfield and van Bommel [(2004). Birds in Europe: Populations estimates, trends and conservation status. Cambridge: BirdLife International]. Overall numbers of bird species maintained by oak and pine archipelagoes were rather similar, but species–area relationships differed between forest types. Intercepts were higher in oak fragments, whereas slopes were steeper in pine plantations. Small oak fragments held more species (mainly Mediterranean Sylvia warblers) than plantations, whereas large plantations held more species than large oak remnants. Differences in species–area relationships seemed to be due to differences in vegetation structure, especially understorey shrub cover and tree height and cover. We recorded nine SPECs, all exclusive (6) or near-exclusive (3) to oak woodlands, although such woodlands do not appear to be critical for their conservation. Hence, we conclude that pine afforestations have played a role for maintaining and restoring forest bird communities in the farming landscapes of central Spain. Promoting large and shrubby plantations would enhance their conservation value for breeding birds, together with promoting growth, regeneration and expansion of Holm oak remnants by means of set-aside measures previous or alternative to oak reafforestation. The increasing importance of non-commercial as compared to commercial values of Mediterranean forests would justify subsidizing the proposed policy.     

Thinning improves growth and resilience after severe droughts in Quercus subpyrenaica coppice forests in the Spanish Pre-Pyrenees

During the past years, growth and productivity of different oak species have been constrained by water shortage in seasonally dry regions such as the Mediterranean Basin. Thinning could improve oak radial growth in these drought-prone regions through the reduction of tree competition for soil water in summer. However, we still lack adequate, long-term assessments on how lasting are thinning treatments effects and to what extent they contribute to oak growth recovery after drought. Here we aim: (i) to study the radial growth sensitivity to drought of Quercus subpyrenaica in the Spanish Pre-Pyrenees, and (ii) to verify if thinning represents a suitable option to enhance growth resistance to drought and post-drought growth recovery. We analysed basal area increment (BAI) trends in the period 1960–2020 of formerly coppiced oak stands thinned in 1984 and compared them with unthinned plots and also with coexisting Scots pine (Pinus sylvestris) growing in thinned plots. We used the Standardized Precipitation Evapotranspiration Index (SPEI) to estimate the severity of droughts and we also assessed climate-growth relationships. Oaks in thinned plots showed higher BAI (369 mm²) than those in unthinned plots (221 mm²). Growth rates remained higher in thinned than in unthinned plots also under severe drought stress. A severe summer drought in 1986 caused abrupt BAI reductions in both oaks (- 40.5%) and pines (- 40.1%). The positive effect of thinning on growth lasted for over 20 years and slightly declined as canopies closed. In the thinned plots, trees with smaller diameter showed the greatest growth release. Oaks in unthinned plots and Scots pine were more sensitive to short-term droughts in terms of growth reduction than oaks in thinned plots, while long term droughts have similar effects on oaks from both thinned and unthinned plots. Oaks were resilient to drought, showing recovery periods lasting from 1 to 2 years in both thinned and unthinned plots. However, intense and prolonged droughts could strongly reverse the expected growth enhancement of thinned plots, and a greater frequency of droughts would limit coppice growth and productivity thus lengthening the rotation periods.     

Phosphorus limitation and competitive capacities of Pinus halepensis and Quercus ilex subsp. rotundifolia on different soils

Aleppo pine (Pinus halepensis) and the evergreen holm oak (Quercus ilex) dominate forest areas of the Mediterranean Basin. Both species regenerate abundantly after fires: pine through seedlings and holm oak through resprouts. Cumulative nutrient losses caused by frequent fires may have decreased soil nutrient availability in such areas. To assess the role of nitrogen and phosphorus as limiting factors for growth of these species during post-fire recovery, a field fertilisation and competition experiment was conducted in a 5-year post-fire shrubland on calcareous soil, where naturally-regenerated saplings of Aleppo pine and resprouts of interior holm oak (Quercus ilex subsp. rotundifolia) coexist. Three years after fertilisation, relative basal area increment was 56% greater in pines fertilised with 250 kg P ha^–1 than in non fertilised ones. N fertilisation had small or no effects. Interactions between N and P fertilisation were not observed. Growth of Aleppo pine only increased with P fertilisation when neighbours were removed. Hence, the negative effect of neighbours on growth was greater when P availability was enhanced by fertilisation. In contrast, holm oak was able to grow more (110%) in response to increased P supply even without neighbour removal. A common garden experiment was then conducted with potted seedlings to investigate whether the suggested higher competitive capacity of holm oak for P held under a range of P amendments on different soils and competitive situations. P fertilisation increased seedling biomass yield of both species. When P availability increased, a negative effect of neighbours on growth was observed for holm oak and in 70 a lesser extent for Aleppo pine. In conclusion, in the field, holm oak resprouts showed higher competitive ability for P uptake compared to Aleppo pine saplings, but in potted seedlings in common garden conditions this trend was not observed. Therefore holm oak is not always competitively superior to Aleppo pine for P. Potted seedlings of both species had a notable plasticity in shoot/root biomass allocation, but only holm oak increased its proportional allocation to roots when neighbours were present. P availability can be a key factor in growth and competitive relations of these two species, but effects differ depending on soil type, individual age, regeneration type (i.e., seedling versus resprouts), and competitive situation. This revised version was published online in June 2006 with corrections to the Cover Date.     

The role of climate change in the widespread mortality of holm oak in open woodlands of Southwestern Spain

Forest decline and increasing tree mortality are of global concern and the identification of the causes is necessary to develop preventive measures. Global warming is an emerging factor responsible for the increasing tree mortality in drought-prone ecosystems. In the southwestern Iberian Peninsula, Mediterranean holm oak open woodlands currently undergo large-scale population-level tree die-off. In this region, temperature and aridity have increased during recent decades, but the possible role of climate change in the current oak mortality has not been investigated.To assess the role of climate change in oak die-off in managed open woodlands in southwestern Spain, we analyzed climate change-related signals in century-long tree ring chronologies of dead holm oaks. We examined the high/low-frequency variability in growth and the relationship between growth and climate.Similar to other Mediterranean forests, growth was favored by precipitation from autumn of the year prior to ring formation to spring of the year of ring formation, whereas high temperatures during spring limited growth. Since the 1970s, the intensity of the high-frequency response to water availability increased simultaneously with temperature and aridity. The growth trends matched those of climatic changes. Growth suppressions occurred during droughts in the 1970s, 1980s and 1990s. Widespread stand-level, age-independent mortality occurred since 2005 and affected trees that cannot be considered old for the species standards.The close relationship between growth and climate indicate that climate change strongly controlled the growth patterns. This suggests that harsher climatic conditions, especially increased aridity, affected the tree performance and could have played a significant role in the mortality process. Climate change may have exacerbated or predisposed trees to the impact of other factors (e.g. intense management and pathogens). These observations could suggest a similar future increase in oak mortality which may occur in more northern oak open woodlands if aridity further increases.     

Effect of cleaning and thinning on height growth and girth increment in holm oak coppices (Quercus ilex L.)

This article studies the influence of cleaning or thinning intensity and of coppice age at the time of silvicultural treatment, on the growth of shoots remaining on the stool, in Quercus ilex coppices.Two experimental designs were used to answer these questions. The first studied the effect of two cleaning intensities on 4, 8, 15, 20, and 25 year old coppices. The second studied the effect of 4 thinning intensities (26%, 42%, 58%, or 78% removal of the initial basal area) on 43 and 57 year old coppices. In both cases, control plots undergoing no silvicultural treatment were maintained for each age class.The results, 4 and 5 years after silvicultural treatment, concerned girth increment and height growth of coppice shoots.Cleaning or thinning consistently had a positive influence on girth increment. In young coppices, less than 25 years old, mean annual girth increment was approximately 6 mm in the controls and double that under heavy cleaning. In older coppices, girth increment was lower in controls (3 mm per year) but the effect of thinning was very positive and, under very heavy thinning, growth was more than 11 mm per year.Cleaning and thinning often modify height growth. This effect was found to be either positive or negative and no general trend could be defined which was valid for all the age classes.Silvicultural treatments also had a positive effect on the appearance and growth of epicormic shoots on the holm oak trunks.The results were also analysed at the stool level. The number or percentage of shoots removed had a positive effect on the individual growth of selected shoots.     

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.     

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.     

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

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

Tree vigour influences secondary growth but not responsiveness to climatic variability in Holm oak

Many tree species from Mediterranean regions have started to show increased rates of crown defoliation, reduced growth, and dieback associated with the increase in temperatures and changes in the frequency and intensity of drought events experienced during the last decades. In this regard, Quercus ilex L. subsp. ballota [Desf.] (Holm oak), despite being a drought-tolerant species widely distributed in the Mediterranean basin, it has recently started to show acute signs of decline, extended areas from Spain being affected. However, few studies have assessed the role of climatic variability (i.e., temperature, precipitation, and drought) on the decline and resilience of Holm oak. Here, we measured secondary growth of seventy Holm oaks from a coppice stand located in central Spain. Sampled trees had different stages of decline, so they were classified into four vigour groups considering their crown foliar lost: healthy (0%), low defoliated (<25%), highly defoliated (25–70%), and dying (70–100%). Our results showed that during the study period (1980–2009) the highly defoliated and dying Holm oaks grew significantly less than their healthy and low defoliated neighbours, suggesting permanent growth reduction in the less vigorous individuals. Despite these differences, all four vigour groups showed similar responses to climatic variations, especially during winter and late spring – early summer seasons, and similar resilience after severe drought events, managing to significantly recover to pre-drought growth rates after only two years. Our findings, hence, illustrate that tree vigour influences secondary growth but not responsiveness to climatic variability in Holm oak. Still, as reduced growth rates are frequently associated with the process of tree mortality, we conclude that the less vigorous Holm oaks might not be able to cope with future water stress conditions, leading to increased mortality rates among this emblematic Mediterranean species.     

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.