Arboreal and prostrate conifers coexisting in Mediterranean high mountains differ in their climatic responses

In contrast to most high elevation areas, plant growth at Mediterranean mountains is exposed to a summer drought period, which represents an additional climatic constraint to low temperatures. Although arboreal and shrubby conifers coexist at high altitudes, most dendroecological studies have focused on climatic responses of tree species, whereas those of shrubby species remain mostly unexplored. We built tree-ring width chronologies for two conifer species, a shrub (Juniperus sabina) and a tree (Pinus sylvestris), coexisting at three high-altitude localities of the Iberian System mountains, eastern Spain. We analyzed their climate–growth relationships for the period 1950–2009 using correlation analyses and multiple regressions. Coexisting species responded to year-to-year climatic variability in different ways. Radial growth in junipers and pines responded positively to April and May temperatures, respectively. Summer drought constrained growth in both cases, although its impact was stronger on junipers than on pines. Our findings suggest that junipers respond earlier than pines to spring temperatures due to their prostrate morphology which may enhance a fast warming of their cambial meristems after snowmelt. The higher dependence of J. sabina on summer rainfall as compared with co-occurring pines confirms that drought stress negatively impacts secondary growth in Mediterranean mountains. This sensitivity to water availability may be caused by the juniper shallow root systems, which mainly use superficial soil water. The climatic signal registered in J. sabina allows studying the response of other similar shrubby woody species growing in Mediterranean alpine areas to the ongoing climate warming, which could also reduce water availability.     

Frugivorous birds mediate sex‐biased facilitation in a dioecious nurse plant

Abstract. Facilitation by dispersal occurs if the nurse plant acts as a focus which is actively selected by seed dispersers and enhances the fitness of the facilitated plant. Sex‐biased facilitation may be produced if seed dispersers tend to concentrate the seeds under female, fruit‐bearing plants of dioecious species more often than under conspecific males. Juniperus sabina is a dioecious shrub with a prostrate growth form from Mediterranean high mountains that modifies many microhabitat characteristics related to seedling establishment and survival. Soil water availability, maximum soil temperature in summer, organic matter and total nitrogen content, were different on open ground as compared with beneath J. sabina shrubs, irrespective of its sex. Other studied characteristics such as soil bulk density and soil compaction after rain did not differ between the microhabitats considered. Some species, such as Juniperus communis, Pinus nigra, Helleborus foetidus and Euphorbia nicaeensis, are spatially associated to J. sabina shrubs, strongly suggesting a facilitative role. The anemochorous P. nigra and myrme‐chorous H. foetidus and E. nicaeensis did not associate preferentially to any sex of J. sabina. Only J. communis, an endozoochorous species sharing the same bird dispersers as J. sabina, presented a female‐biased spatial association with the nurse plant. Seed dispersal mediated by birds attracted by the fruit‐rewarding females of J. sabina explains the sex‐biased spatial pattern of Juniperus communis.     

Climatic factors controlling Pinus sylvestris radial growth along a transect of increasing continentality in southern Siberia

The forest-steppe ecotone in southern Siberia is characterized by a strong dependence of tree growth on summer drought, which is expected to increase under ongoing climate change, with potential consequences for regional and global water and carbon cycles. Since climate conditions control tree secondary growth throughout the growing season, it is assumed that climate change will differently impact the formation of particular tree-ring sectors.In this study, we evaluated spatiotemporal trends in Pinus sylvestris L. tree-ring traits: tree-ring (RW), earlywood (EW) and latewood (LW) widths, as well as their climate response in order to understand potential reactions of P. sylvestris radial growth to climate change along a 4900 km longitudinal transect of increasing continentality in southern Siberia.Results indicated an increasing tree radial growth from the West to the East along the transect. Tree-ring parameters were sensitive to drought, showing a temporal delay in the climatic signals of LW (summer) relative to EW (spring). Climate control of tree growth was stronger at the western site, while it was alleviated over time in eastern sites.This study highlighted the wide plasticity of P. sylvestris to thrive within a wide range of climatic conditions, also suggesting that future drought, as predicted by climate change simulations, will potentially impact P. sylvestris growth differently along the studied gradient, being more susceptible at the western sites due to the current growth limitation.     

Tree species from contrasting hydrological niches show divergent growth and water-use efficiency

Tree species occupy different hydrological niches and climate warming may affect tree performance in those niches through increased drought stress. However, the effects of climate warming on growth, carbon and water fluxes would differ depending on species’ hydrological niche. Species from wet sites should show a lower growth dependence on precipitation and also lower intrinsic water-use efficiency (WUEi), as compared with species from dry sites which should improve more the WUEi. We test these ideas by comparing radial-growth rates (basal-area increment), climate- and drought-growth associations and WUEi of hackberry (Celtis australis) vs. Aleppo pine (Pinus halepensis) and maritime pine (Pinus pinaster) in two Mediterranean sites located in Spain. Species are subjected to similar regional climate conditions in each site but occupy contrasting local hydrological niches (hackberry in wet sites and pines in dry sites). Climate is warming in both study sites and drought-avoiding pines are responding by showing higher growth rates and improved WUEi. We also found a similar growth dependency on winter-spring precipitation and summer drought of all species and sites and comparable WUEi values and trends, excepting in hackberry from southern Spain which grew more, and showed a higher growth resistance to drought and lower and more stable WUEi values. Variables inferred from tree rings as growth rates and WUEi allow characterizing the hydrological niche of tree species, which may be contingent on site conditions and climate warming.     

Earlier summer drought affects leaf functioning of the Mediterranean species Cistus monspeliensis L.

The Mediterranean vegetation is characterized by a high diversity of growth forms, habits and phenology that enable it to endure under harsh environmental conditions. It is however unclear whether these adaptations may allow plant survival under more extreme conditions, as predicted by climatic models under the perspective of climate change. A manipulative experiment aiming at anticipating summer aridity has been run to analyse the effects of the experimental drought on spring-leaf functioning and characteristics of the leaf-dimorphic Mediterranean shrub Cistus monspeliensis L.Assimilation rates were reduced under anticipated summer aridity due to a decrease of stomatal conductance, but only before morphological adaptations to drought (an increase of leaf mass per area) occurred. These adaptations were anticipated under experimental dry conditions, and causes photosynthetic performances to recover compared to previous dates. When natural summer aridity occurred, the leaf mass per area also changed in the control. However, this causes no recovery of the photosynthetic performances, because of the decrease of stomatal conductance due to low soil water content and leaf water potential values. Moreover, under experimental drought, leaf shedding was anticipated to reduce water losses, causing an overall reduction of leaf lifespan.     

Growth and carbon isotopes of Mediterranean trees reveal contrasting responses to increased carbon dioxide and drought

Forest dynamics will depend upon the physiological performance of individual tree species under more stressful conditions caused by climate change. In order to compare the idiosyncratic responses of Mediterranean tree species (Quercus faginea, Pinus nigra, Juniperus thurifera) coexisting in forests of central Spain, we evaluated the temporal changes in secondary growth (basal area increment; BAI) and intrinsic water-use efficiency (iWUE) during the last four decades, determined how coexisting species are responding to increases in atmospheric CO₂ concentrations (C _a) and drought stress, and assessed the relationship among iWUE and growth during climatically contrasting years. All species increased their iWUE (ca. +15 to +21 %) between the 1970s and the 2000s. This increase was positively related to C _a for J. thurifera and to higher C _a and drought for Q. faginea and P. nigra. During climatically favourable years the study species either increased or maintained their growth at rising iWUE, suggesting a higher CO₂ uptake. However, during unfavourable climatic years Q. faginea and especially P. nigra showed sharp declines in growth at enhanced iWUE, likely caused by a reduced stomatal conductance to save water under stressful dry conditions. In contrast, J. thurifera showed enhanced growth also during unfavourable years at increased iWUE, denoting a beneficial effect of C _a even under climatically harsh conditions. Our results reveal significant inter-specific differences in growth driven by alternative physiological responses to increasing drought stress. Thus, forest composition in the Mediterranean region might be altered due to contrasting capacities of coexisting tree species to withstand increasingly stressful conditions.     

Silver birch shows nonlinear responses to moisture availability and temperature in the eastern Baltic Sea region

Silver birch (Betula pendula Roth.) is a widespread species with a high potential for aiding sustainability and multifunctionality of European forests, as evidenced in Finland and the Baltics. However, under increasing relevance of climate change for tree growth, the meteorological sensitivity of the species is largely unknown, presuming it to be weather tolerant (low sensitivity). Considering local adaptations of populations of widespread species, climatic changes are subjecting trees to extreme conditions, thus testing their adaptability. Accordingly, information on the plasticity (variability) of responses across a gradient of meteorological conditions is crucial for reliable predictions of tree growth. Tree-ring width network was established to assess the plasticity of growth responses of silver birch to meteorological conditions across the eastern Baltic climatic gradient. Time series analysis in combination with generalized additive modelling were applied to assess responses of birch from 21 naturally regenerated conventionally managed stands scattered from southern Finland to northern Germany. Despite the presumed tolerance, explicit meteorological sensitivity of silver birch was estimated. A gradient of local linear weather-growth relationships was estimated, as growth limitation shifted from temperature during the dormancy to water availability during vegetation period in southern Finland and northern Germany, respectively. However, these relationships were nonstationary, as the effect of summer water shortage was intensifying and sensitivity to it has likely been subjected to local adaptation. The regional generalization revealed presence of stationary, yet nonlinear and plastic growth responses, implying disproportional effects of climatic changes. Such responses also explained the nonstationarities, as the local climates shifted along the regional gradient. At the regional scale, summer water shortage was the main driver of increment, while winter conditions had a secondary role; temperature of the preceding vegetation season also had an effect on increment. Accordingly, increased variability of increment of silver birch is expected under changing climate; still, sensitivity and plasticity of increment can be considered as an adaptation to shifting environments.     

Plastic bimodal growth in a Mediterranean mixed-forest of Quercus ilex and Pinus halepensis

Mediterranean tree species have evolved to face seasonal water shortages, but may fail to cope with future increases in drought frequency and intensity. We investigated stem radial increment dynamics in two typical Mediterranean tree species, Aleppo pine (Pinus halepensis), a drought-avoiding species, and holm oak (Quercus ilex), a drought-tolerant species, in a mixed forest and on contrasting slope aspects (south- and north-facing). Intra- and inter-annual growth patterns were modelled using the VS-Lite2 model for each tree species and slope-aspect. Both species showed a bimodal growth pattern, with peaks coinciding with favourable conditions in spring and autumn. A bimodal growth pattern is always observed in P. halepensis, while in Q. ilex is facultative, which suggests different strategies adopted by these species to cope with summer drought. More specifically, trees on south-facing slope showed a more evident bimodal pattern and more intra-annual density fluctuations. In recent decades, the intensity of both growth peaks has diminished and drifted away due to the increased summer drought. The VS-Lite2 model reveals a niche partitioning between both species. Differences in growing season’s length and timings of growth peaks in both species are relevant for their coexistence and should be considered for estimating mixed-forest responses under climate change scenarios.     

Climatic signal in growth-rings of Copaifera lucens: An endemic species of a Brazilian Atlantic forest hotspot,southeastern Brazil

In this study, we present the first tree-ring chronology for the tropical tree species Copaifera lucens and its climatic signal in southeastern Brazil. Tree-ring width series were compared with local climate indices using a drought index (Standardized Precipitation Evapotranspiration Index —SPEI), in monthly, bi-monthly and four-monthly scales. We also calculated negative pointer years over the time-span of the tree-ring width. The radial growth of C. lucens showed a positive correlation with the SPEI of the current summer and autumn in all the three analyzed time scales, while the negative pointer years matched with drier years. The species was highly sensitive to very low summer precipitation, which may lead to a 49% reduction in growth. We conclude that the long-living C. lucens has a great potential for dendrochronological studies as it shows a marked climatic signal. Our study also reinforces the importance of rainfall in regulating radial growth in tropical forests and sheds light on the local climate influence on tree growth in recent decades.     

Spatiotemporal variability of stone pine (Pinus pinea L.) growth response to climate across the Iberian Peninsula

Climate warming and increasing aridity have impacted diverse ecosystems in the Mediterranean region since at least the 1970s. Pinus pinea L. has significant environmental and socio-economic importance for the Iberian Peninsula, so a detailed understanding of its response to climate change is necessary to predict its status under future climatic conditions. However, variability of climate and uncertainties in dendroclimatological approach complicate the understanding of forest growth dynamics. We use an ensemble approach to analyze growth-climate responses of P. pinea trees from five sites along a latitudinal gradient in Spain over time. The growth responses to April-June precipitation totals were stronger in the north than in the south. Since the 1950s, the sensitivity of growth to April-June precipitation increased in the north and decreased in the south. Meteorological drought usually started in May in the southern sites, but in June-July in the northern sites. The water deficit in the southern sites is thus greater and more limiting for tree growth, and this likely accounts for the lower growth sensitivity during these months. Our results indicate that P. pinea has a high degree of plasticity, suggesting the species will withstand changing climatic conditions. However, growth response to drought regimes varies among P. pinea populations, suggesting that different populations have different capacities for acclimation to warmer and drier climate, and this may influence future vegetation composition.     

Growth response of Abies georgei to climate increases with elevation in the central Hengduan Mountains,southwestern China

Climatic harshness is expected to increase at higher elevations; however, elevational trends of tree radial growth response of high-elevation forests to climate change need to be investigated at different locations because of existing local variability in site-specific climatic conditions. We developed tree-ring width chronologies of Yunnan fir (Abies georgei) along elevation gradients at two sites in the central Hengduan Mountains (HM). High-elevation forests of A. georgei showed growth synchronicity and common growth signals along elevation gradients, indicating a common climatic forcing, although tree radial growth rates decreased with increasing elevation. Radial growth of Yunnan fir showed positive correlations with summer temperatures and February precipitation and moisture availability, but were negatively correlated with spring temperatures. The strongest positive relationship indicated summer (July) mean and minimum temperatures are the most important growth determining climatic factors for tree radial growth in the cold environment of HM, and this relationship revealed a clear elevational trend with stronger correlations at higher altitudes. In contrast, tree radial growth was negatively correlated with June precipitation and moisture availability. The whole study period 1954–2015 was split in two sub-periods of equal length. Comparing the early sub-period (1954–1984) to the later sub-period (1985–2015), tree growth response to the summer temperatures strongly increased, while it became weaker to June precipitation and moisture availability. High-elevation Yunnan fir forests in the HM currently benefit from elevated growing season temperatures under humid summer conditions. However, increasing temperatures may induce drought stress on tree radial growth if the observed decreasing trend in humidity and precipitation continues.     

Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40‐year tree ring record and a 30‐year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (?10 days) due to winter warming and earlier growth cessation (?26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving‐window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate–growth correlations matches the start of the recent atmospheric warming pause also known as the ‘climate hiatus’. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.     

Growth phenological variations in the narrow-leaved ash (Fraxinus angustifolia) over the Mediterranean region: A simulation study

Tree species inhabiting riparian forests under Mediterranean climate have evolved to face summer water shortage but may fail to cope with future increases in drought severity. Thus, understanding tree growth phenological variations in response to environmental conditions is necessary to assess the impact of seasonal drought in riparian forests. In this study, we investigated the response of stem radial growth to climate in the narrow-leaved ash (Fraxinus angustifolia) over its distribution in southern Europe. We simulated intra- and inter-annual growth patterns using the Vaganov-Shashkin (VS) model considering five sites subjected to summer drought but showing different climate conditions. The growth pattern in this species varied from unimodal in cool-wet sites to facultative bimodal in warm-dry sites. Bimodal patterns were characterized by two growth peaks coinciding with favorable climate conditions in spring and autumn. The spring growth peak occurs earlier (May) in warm-dry sites than in wet-cool sites (June–July). The variation in the season growth length and growth timing suggests different strategies adopted by this species to cope with summer drought. The VS model revealed different growth patterns across which would be relevant in predicting the response of this and other riparian tree species to climate warming and aridification. Differences in the length of the growing season, timings of growth peaks and the shift from unimodal to bimodal growth patterns should be considered when assessing growth adjustments to future climate scenarios.     

Site-contingent responses to drought of core and relict Tetraclinis articulata populations from Morocco and Spain

The growth responses to climate variability are still unknown in locally threatened conifers from dry regions, but this information is necessary for improving the conservation of relict populations under increasing aridification. We characterized the radial growth patterns and responses to climate of Tetraclinis articulata, a Cupressaceae tree endemic to the western Mediterranean Basin, in a relict population located in southwestern Spain (Doñana) and two populations from the northern Morocco where the species core habitat is found (Tétouan, Ifarten). We assessed climate-growth relationships by using tree-ring width, climate data, drought and North Atlantic Oscillation (NAO) indices. Climate-growth analyses were refined using the climwin R package to select the most informative statistical models. The main climatic constraints of growth were inferred by using the process-based Vaganov-Shashkin (VS) model explicitly considering non-linear climate-growth relationships. Tetraclinis articulata growth was favored by wet conditions from the prior autumn to the spring of the growth year. In Doñana, warmer May conditions led to growth decline but this negative effect could be reversed by wet-warm conditions in the prior late autumn. Growth in the two Moroccan sites was constrained by 6- to 18-month long droughts peaking in summer, which account for cumulative water deficit since the previous autumn. Winter and early spring precipitation were the main climate drivers of growth in the Moroccan T. articulata populations, and their year-to-year variability was linked to the NAO. The VS model simulations showed that T. articulata growth is enhanced by wet soil conditions in late winter and early spring, probably recharging shallow soil water pools. The VS model also confirmed that warmer spring-summer conditions could amplify drought stress and threaten the long-term persistence of the relict Doñana population.     

Summer drought exposure,stand structure,and soil properties jointly control the growth of European beech along a steep precipitation gradient in northern Germany

Increasing exposure to climate warming-related drought and heat threatens forest vitality in many regions on earth, with the trees' vulnerability likely depending on local climatic aridity, recent climate trends, edaphic conditions, and the drought acclimatization and adaptation of populations. Studies exploring tree species' vulnerability to climate change often have a local focus or model the species' entire distribution range, which hampers the separation of climatic and edaphic drivers of drought and heat vulnerability. We compared recent radial growth trends and the sensitivity of growth to drought and heat in central populations of a widespread and naturally dominant tree species in Europe, European beech (Fagus sylvatica), at 30 forest sites across a steep precipitation gradient (500–850 mm year⁻¹) of short length to assess the species' adaptive potential. Size-standardized basal area increment remained more constant during the period of accelerated warming since the early 1980s in populations with >360 mm growing season precipitation (April–September), while growth trends were negative at sites with <360 mm. Climatic drought in June appeared as the most influential climatic factor affecting radial growth, with a stronger effect at drier sites. A decadal decrease in the climatic water balance of the summer was identified as the most important factor leading to growth decline, which is amplified by higher stem densities. Inter-annual growth variability has increased since the early 1980s, and variability is generally higher at drier and sandier sites. Similarly, within-population growth synchrony is higher at sandier sites and has increased with a decrease in the June climatic water balance. Our results caution against predicting the drought vulnerability of trees solely from climate projections, as soil properties emerged as an important modulating factor. We conclude that beech is facing recent growth decline at drier sites in the centre of its distribution range, driven by climate change-related climate aridification.     

Growth of 19 conifer species is highly sensitive to winter warming,spring frost and summer drought

Background and AimsConifers are key components of many temperate and boreal forests and are important for forestry, but species differences in stem growth responses to climate are still poorly understood and may hinder effective management of these forests in a warmer and drier future.MethodsWe studied 19 Northern Hemisphere conifer species planted in a 50-year-old common garden experiment in the Netherlands to (1) assess the effect of temporal dynamics in climate on stem growth, (2) test for a possible positive relationship between the growth potential and climatic growth sensitivity across species, and (3) evaluate the extent to which stem growth is controlled by phylogeny.Key resultsEighty-nine per cent of the species showed a significant reduction in stem growth to summer drought, 37 % responded negatively to spring frost and 32 % responded positively to higher winter temperatures. Species differed largely in their growth sensitivity to climatic variation and showed, for example, a four-fold difference in growth reduction to summer drought. Remarkably, we did not find a positive relationship between productivity and climatic sensitivity, but instead observed that some species combined a low growth sensitivity to summer drought with high growth potential. Both growth sensitivity to climate and growth potential were partly phylogenetically controlled.ConclusionsA warmer and drier future climate is likely to reduce the productivity of most conifer species. We did not find a relationship between growth potential and growth sensitivity to climate; instead, some species combined high growth potential with low sensitivity to summer drought. This may help forest managers to select productive species that are able to cope with a warmer and drier future.     

Effect of ecological factors on intra-annual stem girth increment of holm oak

Key message

The intra-annual stem girth increment of Quercus ilex is mainly driven by water availability and secondly by temperature. Tree size and competition modulate the growth response to climate.

Abstract

Holm oak (Quercus ilex ssp. ballota [Desf.] Samp.) is the most widespread species in the Iberian peninsula, being one of the most representative trees in forests and open woodlands. The analysis of stem girth increment of holm oak may provide valuable information about how Mediterranean ecosystems will respond to the forecasted climate changes. However, due to the variability of the Mediterranean climate, the knowledge of intra-annual patterns of growth is needed for a better understanding of the influence of the climatic variables at this scale. To this end, we used band dendrometers to measure monthly stem girth increments of 96 holm oak trees from 2003 to 2010, located in open woodlands and dense Mediterranean forests in southwestern Spain. We assessed the effects of climate, competition, topography, and initial stem diameter on stem girth increment. The major stem increment periods were in spring and autumn whereas increment rates were very low or even negative in winter and summer. Spring was not every year the season with the higher stem increments, but autumn when spring was very dry. Higher precipitation, soil moisture, and relative humidity had significant positive effects on stem increment, whereas higher temperature, reference evapotranspiration, and solar radiation had significant negative effects. Initial tree diameter and competition from nearby trees partly explained significant differences in stem increment of individual trees. Therefore, the forecasted climatic changes, in which decreased rainfall in spring and increased summer drought are expected in the Mediterranean region, may be a significant threat to the Q. ilex ecosystems.     

Winter Climate Limits Subantarctic Low Forest Growth and Establishment

Campbell Island, an isolated island 600 km south of New Zealand mainland (52°S, 169°E) is oceanic (Conrad Index of Continentality  = −5) with small differences between mean summer and winter temperatures. Previous work established the unexpected result that a mean annual climate warming of c. 0.6°C since the 1940''s has not led to upward movement of the forest limit. Here we explore the relative importance of summer and winter climatic conditions on growth and age-class structure of the treeline forming species, Dracophyllum longifolium and Dracophyllum scoparium over the second half of the 20^th century. The relationship between climate and growth and establishment were evaluated using standard dendroecological methods and local climate data from a meteorological station on the island. Growth and establishment were correlated against climate variables and further evaluated within hierarchical regression models to take into account the effect of plot level variables. Winter climatic conditions exerted a greater effect on growth and establishment than summer climatic conditions. Establishment is maximized under warm (mean winter temperatures >7 °C), dry winters (total winter precipitation <400 mm). Growth, on the other hand, is adversely affected by wide winter temperature ranges and increased rainfall. The contrasting effect of winter warmth on growth and establishment suggests that winter temperature affects growth and establishment through differing mechanisms. We propose that milder winters enhance survival of seedlings and, therefore, recruitment, but increases metabolic stress on established plants, resulting in lower growth rates. Future winter warming may therefore have complex effects on plant growth and establishment globally.     

Reproductive output in Mediterranean shrubs under climate change experimentally induced by drought and warming

The effects of climate change on plant reproductive performance affects the sequence of different plant reproductive stages from flowering to seed production and viability, as well as the network of relationships between them. These effects are expected to respond to different components of climate change, such as temperature and water availability, and may be sensitive to differences in species phenology.We used long-term experimental drought and warming treatments to study the effect of climate change on flower production, fruit and seed-set, seed size and seed germination rate (proportion of germinating seeds) in three Mediterranean shrubs coexisting in a coastal shrubland.Larger plants produced significantly more flowers in all three species, and higher fruit-set in Dorycnium pentaphyllum. Flower production was reduced in drought and warming treatments in the spring-flowering species D. pentaphyllum and Helianthemum syriacum, but not in the autumn–winter species Erica multiflora, which increased flowering in the warming treatment. However, the drought treatment eventually resulted in a decreased seed-set in E. multiflora. Structural equation modelling revealed strong correlations between the sequential reproductive stages. Specifically, flower density in inflorescences determined seed-set in H. syriacum, and seed size and germination rate in E. multiflora. Nevertheless, the relevance of relationships between reproductive traits changed between climatic treatments: in D. pentaphyllum a direct relationship between plant size and seed size only arised in the drought treatment, while in H. syriacum climate treatments resulted in a stronger relationship between the number of flowers and seed-set.This experimental study shows the ability of changing climatic variables to determine the reproductive sequential process of woody species. We show that several parameters of the reproductive performance of some Mediterranean species are affected by drought and warming treatments simulating climate change, highlighting the importance of changes in both water availability and temperature, and the sequential relationship between reproductive stages. Phenological patterns also contribute to species’ differential responses to climatic change, due to the relationship of these patterns with resource availability, environmental conditions and plant–pollinator interactions.     

Miocene to Pliocene vegetation reconstruction and climate estimates in the Iberian Peninsula from pollen data

Pollen analysis of Miocene and Pliocene sediments from the Iberian Peninsula shows a progressive reduction in plant diversity through time caused by the disappearance of thermophilous and high-water requirement plants. In addition, an increase in warm-temperate (mesothermic), seasonal-adapted “Mediterranean” taxa, high-elevation conifers and herbs (mainly Artemisia) occurred during the Middle and Late Miocene and Pliocene. This has mainly been interpreted as a response of the vegetation to global and regional processes, including climate cooling related to the development of the East Antarctic Ice Sheet and then the onset of the Arctic Ice Sheet, uplift of regional mountains related to the Alpine uplift and the progressive movement of Eurasia towards northern latitudes as a result of the northwards subduction of Africa. The development of steppe-like vegetation in southern Iberia is ancient and probably started during the Oligocene. The onset of a contrasted seasonality in temperature during the Mid-Pliocene superimposed on the pre-existing seasonality in precipitation, the annual length of which increased southward. The Mediterranean climatic rhythm (summer drought) began about 3.4 Ma and caused the individualization of modern Mediterranean ecosystems. Quaternary-type Mediterranean climatic fluctuations started at 2.6 Ma (Gelasian) resulting in repeated steppe vs. forest alternations. A latitudinal climatic gradient between the southern and the northern parts of the Iberian Peninsula existed since the Middle Miocene.