Hydraulic properties and freezing-induced cavitation in sympatric evergreen and deciduous oaks with contrasting habitats

We investigated the hydraulic properties in relation to soil moisture, leaf habit, and phylogenetic lineage of 17 species of oaks (Quercus) that occur sympatrically in northern central Florida (USA). Leaf area per shoot increased and Huber values (ratio of sapwood area to leaf area) decreased with increasing soil moisture of species’ habitats. As a result, maximum hydraulic conductance and maximum transpiration were positively correlated with mean soil moisture when calculated on a sapwood area basis, but not when calculated on a leaf area basis. This reveals the important role that changes in allometry among closely related species can play in co‐ordinating water transport capacity with soil water availability. There were significant differences in specific conductivity between species, but these differences were not explained by leaf habit or by evolutionary lineage. However, white oaks had significantly smaller average vessel diameters than red oaks or live oaks. Due to their lower Huber values, maximum leaf specific conductivity (K_L) was higher in evergreen species than in deciduous species and higher in live oaks than in red oaks or white oaks. There were large differences between species and between evolutionary lineages in freeze–thaw‐induced embolism. Deciduous species, on average, showed greater vulnerability to freezing than evergreen species. This result is strongly influenced by evolutionary lineage. Specifically, white oaks, which are all deciduous, had significantly higher vulnerability to freezing than live oaks (all evergreen) and red oaks, which include both evergreen and deciduous species. These results highlight the importance of taking evolutionary lineage into account in comparative physiological studies.     

Summer and winter sensitivity of leaves and xylem to minimum freezing temperatures: a comparison of co-occurring Mediterranean oaks that differ in leaf lifespan

Freezing sensitivity of leaves and xylem was examined in four co-occurring Mediterranean oaks (Quercus spp.) grown in a common garden to determine whether freezing responses of leaves and xylem were coordinated and could be predicted by leaf lifespan. Freezing-induced embolism and loss of photosynthetic function were measured after overnight exposure to a range of subzero temperatures in both summer and winter. Both measures were found to be dependent on minimum freezing temperature and were correlated with leaf lifespan and vessel diameter. The dependence of xylem embolism on minimum freezing temperature may result from the decline in water potential with ice temperature that influences the redistribution of water during freezing and leads to an increase in xylem tension. Winter acclimatization had a relatively small effect on the vulnerability to freezing-induced embolism, although leaf photosynthetic function showed a strong acclimatization response, particularly in the two evergreen species. Quercus ilex, the species with the longest leaf lifespan and narrowest vessel diameters, showed the highest freezing tolerance. This helps explain its ability to inhabit a broad range throughout the Mediterranean region. By contrast, the inability of the deciduous oaks to maintain photosynthetic and vascular function throughout the winter indicates a competitive disadvantage that may prevent them from expanding their ranges.     

Predicting vegetation type through physiological and environmental interactions with leaf traits: evergreen and deciduous forests in an earth system modeling framework

Earth system models are incorporating plant trait diversity into their land components to better predict vegetation dynamics in a changing climate. However, extant plant trait distributions will not allow extrapolations to novel community assemblages in future climates, which will require a mechanistic understanding of the trade‐offs that determine trait diversity. In this study, we show how physiological trade‐offs involving leaf mass per unit area (LMA), leaf lifespan, leaf nitrogen, and leaf respiration may explain the distribution patterns of evergreen and deciduous trees in the temperate and boreal zones based on (1) an evolutionary analysis of a simple mathematical model and (2) simulation experiments of an individual‐based dynamic vegetation model (i.e., LM3‐PPA). The evolutionary analysis shows that these leaf traits set up a trade‐off between carbon‐ and nitrogen‐use efficiency at the scale of individual trees and therefore determine competitively dominant leaf strategies. As soil nitrogen availability increases, the dominant leaf strategy switches from one that is high in nitrogen‐use efficiency to one that is high in carbon‐use efficiency or, equivalently, from high‐LMA/long‐lived leaves (i.e., evergreen) to low‐LMA/short‐lived leaves (i.e., deciduous). In a region of intermediate soil nitrogen availability, the dominant leaf strategy may be either deciduous or evergreen depending on the initial conditions of plant trait abundance (i.e., founder controlled) due to feedbacks of leaf traits on soil nitrogen mineralization through litter quality. Simulated successional patterns by LM3‐PPA from the leaf physiological trade‐offs are consistent with observed successional dynamics of evergreen and deciduous forests at three sites spanning the temperate to boreal zones.     

Restoring for the present or restoring for the future: enhanced performance of two sympatric oaks (Quercus ilex and Quercus pyrenaica) above the current forest limit

Reforestation is common to restore degraded ecosystems, but tree‐species choice often neglects ongoing environmental changes. We evaluated the performance of planted seedlings of two oak species at two sites in a Mediterranean mountain (Sierra Nevada, SE Spain): one located within the current altitudinal forest range (1,600–1,760 m), and one above the upper forest limit (1,970–2,120 m). The forest service planted 1,350 seedlings of the deciduous Pyrenean oak and the evergreen Holm oak in a postfire successional shrubland. After 2 years, seedlings were monitored for survival, and a subset of 110 Pyrenean oaks and 185 Holm oaks were harvested for analyses of biomass and foliar nutrient status, δ¹³C, and δ¹⁸O. Both species showed the highest survival and leaf N status above the upper forest limit, and survival increased with altitude within each plot. The deciduous oak benefited most from planting at higher altitude, and it also had greater biomass at the higher site. Correlations between foliar N, δ¹⁸O, and δ¹³C across elevations indicate tighter stomatal control of water loss and greater water‐use efficiency with increasing plant N status at higher altitude, which may represent a so‐far overlooked positive feedback mechanism that could foster uphill range shifts in water‐limited mountain regions. Given ongoing trends and future projections of increasing temperature and aridity throughout the Mediterranean region, tree‐species selection for forest restoration should target forecasted climatic conditions rather than those prevailing in the past. This study highlights that ecosystem restoration provides an opportunity to assist species range shifts under rapidly changing climate.     

壳斗科的地质历史及其系统学和植物地理学意义

在收集整理现有壳斗科化石资料的基础上,讨论了壳斗科及其各属的起源时间、地史分布和地史 演替过程以及这些化石资料在系统学和植物地理学上的意义。白垩纪尚无壳斗科可靠的大化石记录, 微化石需要进一步研究才能确定亲缘关系以及古新世壳斗科已经分化出两个类群。从以上这些事实推 论壳斗科起源于白垩纪晚期,而壳斗科现代各属出现的时间应不晚于古新世。最早发现的壳斗科化石和现代栗亚科和水青冈亚科在形态结构上非常相似,这一事实表明,壳斗科分为两个亚科的观点更接近客观事实。在水青冈亚科中,三棱栎类的化石最早出现;在栎属中,青冈亚属更接近祖先类群;在地史中全缘栎类较具齿栎类出现早,粗齿的落叶栎类出现最晚。三棱栎属、栲属和石栎属的化石在老第三纪出现于北美和欧洲的事实说明,北美、欧洲和东亚在老第三纪时有一个相通的壳斗科植物区系。南美的三棱栎是通过北美进入南美的。中国横断山、欧洲地中海沿岸和北美西北部有一类形态特征相似、亲缘关系相近的硬叶栎类,它们之间有相同的地质演替历史,它们现代分布边界可能就是古地中海的边界。美洲的栎类有两个来源,常绿硬叶栎类是通过古地中海沿岸而经北美-欧洲陆桥到达的,落叶栎类则是在中新世以后通过白令海峡到达的。     

On the relationship between abiotic stress and co-occurrence patterns: an assessment at the community level using soil lichen communities and multiple stress gradients

The stress‐gradient hypothesis (SGH) predicts a shift from predominant competition to facilitation as abiotic stress increases. Most empirical tests of the SGH have evaluated the interactions between a single or a few pairs of species, have not considered the effects of multiple stress factors, and have not explored these interactions at nested spatial scales. We sampled 63 0.25‐m² plots, each subdivided into 100 5×5 cm and 25 10×10 cm sampling squares, in a semi‐arid Mediterranean environment to evaluate how co‐occurrence patterns among biological soil crusts (BSC)‐forming lichens changed along natural stress gradients driven by water and nutrient (N, P, K) availability. According to the SGH, we tested the hypothesis that the fine‐scale spatial arrangement of BSC‐forming lichens should shift from prevailing interspecific segregation to aggregation as abiotic stress increases. Co‐occurrence patterns ranged from significant species segregation to aggregation at the two spatial scales studied. When using the 5×5 cm grid, more plots showed significant species segregation than aggregation. At this sampling scale, co‐occurrence increased as water and nutrient availability decreased and increased, respectively. Small increases in soil pH promoted species co‐occurrence. Interspecific segregation was promoted as the cover of highly competitive species, such as Diploschistes diacapsis, increased. No significant relationships between co‐occurrence and the surrogates of abiotic stress were observed when data was arranged in a 10×10 cm grid. Our co‐occurrence analyses partially supported predictions from the SGH, albeit the results obtained were dependent on the type of abiotic stress and the spatial scale considered. They show the difficulties of predicting how co‐occurrence patterns change along complex stress gradients, and highlight the need of incorporating the effects of abiotic stress promoted by different resources, such as water and nutrients, into the conceptual framework of the SGH.     

鄂西南亚热带山地常绿落叶阔叶混交林物种多度分布格局

基于七姊妹山自然保护区内6 hm~2监测样地多度数据,通过累计经验分布曲线(ECDF)表征该样地内不同生活型功能群的物种-多度分布格局,并采用6种模型对各功能群不同取样尺度物种等级-多度曲线进行拟合并检验其拟合效果,分析多度格局与模型拟合在不同尺度间的差异,探讨其背后的生态学过程与机制。结果表明:(1)各尺度下落叶种比常绿种的物种数多,物种多样性指数更大,但个体数相对较少;不同功能群稀有种比例排序为:落叶种所有种常绿种。(2)6种模型中的断棍模型的拟合效果较差;中大尺度(50 m×50 m、100 m×100 m)上不同生活型树种多度分布能接受的模型较少,除大尺度的常绿树种外,拟合最优模型均为对数正态分布模型,大尺度的常绿树种拟合最优模型为中性模型;小尺度上(20 m×20 m)常绿树种的最优模型为对数正态分布模型,落叶树种最优模型为生态位优先模型,所有树种在小尺度最优模型为Zipf-Mandelbrot模型。研究认为,随着尺度逐渐扩大,中性过程较生态位过程对物种-多度格局的解释力度更大,落叶树种物种多度格局的形成机制较常绿树种更接近于样地所有树种物种-多度格局的形成机制。     

Comparative ecophysiological responses to drought of two shrub and four tree species from karst habitats of southwestern China

Drought stress is one of the most important factors in limiting the survival and growth of plants in the harsh karst habitats of southwestern China, especially at the seedling establishment stage. The ecophysiological response to drought stress of native plants with different growth forms is useful for re-vegetation programs. Two shrub and four tree species were studied, including Pyracantha fortuneana (evergreen shrub), Rosa cymosa (deciduous shrub), Cinnamomum bodinieri (evergreen tree), and other three deciduous trees, Broussonetia papyrifera, Platycarya longipes, and Pteroceltis tatarinowii. The seedlings were randomly assigned to four drought treatments, i.e., well-watered, mild drought stress, moderate drought stress, and severe drought stress. Leaf water relations, gas exchange, chlorophyll fluorescence, and growth of the seedlings were investigated. Under severe drought stress, the two shrubs with low leaf area ratio (LAR) maintained higher water status, higher photosynthetic capacity, and larger percent biomass increase than the most of the trees. The two shrubs also had lower specific leaf area, greater intrinsic water use efficiency, and thermal dissipation than the trees. This suggested that the two shrubs had high tolerance to severe drought and were suitable for re-vegetation in harsh habitats. The evergreen C. bodinieri exhibited higher leaf mass ratio (LMR) and LAR than the deciduous species under mild and moderate stress. However, the low maximum quantum efficiency of PSII photochemistry (F _v/F _m) and net assimilation rate, and the sharp decreases of water potential, LMR, LAR, and biomass under severe stress indicated C. bodinieri’s weak tolerance to severe drought. In response to drought stress, the three deciduous trees revealed sharp reductions of biomass due to the large drought-induced decreases of gas exchange, LAR, and LMR. Under drought conditions, the deciduous trees minimized water loss by stomatal closure and by reducing transpiration leaf area and light harvesting through shedding leaves. This suggested that the three deciduous trees were more sensitive to water availability than the shrubs and used avoidance strategies against drought stress. However, the better growth performance of the deciduous trees than that of the shrubs under favorable conditions suggested that deciduous trees could be suitable for habitats with mild and temporary drought stress.     

Response of broadleaved evergreen Mediterranean forest vegetation to fire disturbance during the Holocene: insights from the peri-Adriatic region

Aim To test whether fire contributed to the expansion and compositional change of evergreen forests in the Mediterranean region during the Holocene. Location The peri‐Adriatic region, encompassing the Italian peninsula, Sicily and the western and southern Balkans between latitudes 46° and 37° N. Methods New high‐resolution pollen and microscopic charcoal data from Lago dell’Accesa (Tuscany, Italy) were used to estimate the response of the evergreen oak, Quercus ilex L., to fire during its expansion phase at 8500 cal. yr bp . The data were compared with the pollen and charcoal series from other Mediterranean sites (Lago di Massaciuccoli in Tuscany, Malo Jezero in Croatia, Biviere di Gela in Sicily) and analysed using numerical techniques (redundancy analysis, detrended canonical correspondence analysis) to identify long‐term fire–vegetation linkages and the degree of compositional change. Results Microscopic charcoal and pollen of evergreen oaks were negatively correlated during the period of quasi‐natural fire regime (Mesolithic, 10,000–8000 cal. yr bp ). In addition, there was no such positive correlation during periods when the fire regime was potentially more influenced by people (Neolithic–Bronze Age, 8000–3000 cal. yr bp ). Compared with inland sites, coastal sites that are currently located at a distinct ecotone showed more compositional change. Main conclusions The analyses suggest that climatic change, without an additional effect of fire regimes, favoured the expansion and compositional change of evergreen forests across the peri‐Adriatic region. Strikingly different patterns occurred along a north–south gradient. In the north (Tuscany and Croatia, meso‐Mediterranean belt), Q. ilex replaced deciduous forests when conditions became drier; in the south (Sicily, thermo‐Mediterranean belt) the species replaced maquis or steppe vegetation when climatic conditions became moister. We conclude that the projected increase in fire activity may lead to the loss of most of the remaining relict forests of Q. ilex in southern Europe.     

Sculpture of pollen grains of <Emphasis Type="Italic">Quercus</Emphasis> L. from the holocene of the south of the Sea of Japan

Fossil pollen grains of Quercus L. were studied with application of an EVO 40 scanning electron microscope. The material came from Holocene core samples from deep water zone of the south of the Sea of Japan and shelf zone of East Korea Bay. The pollen grains were obtained from three radiocarbon-dated palynological assemblages. Six types of fossil pollen grains were revealed: four of them were assigned to deciduous oaks and two were assigned to evergreen oaks.     

Comparative water relations of four Mediterranean oak species

The water relations and responses of two evergreen (Quercus ilex L. and Q. suber L.) and two deciduous (S. afares Pomel. and Q. faginea Will.) Quercus species were studied under experimental conditions. Two-year old seedlings grown in 30 l. pots were subjected to a drying period during which stomatal conductance, pre-dawn potential and minimum foliar potential were measured.The results shows that, for all species, the daily course of stomatal conductance agrees with the patterns proposed by Hinckley et al. (1978 & 1983). Concurrent with the species responses to short-term variation in water availability, it was found that pre-dawn leaf water potential controlled the maximum daily leaf conductance. There was a strong correlation between pre-dawn leaf potential and maximum daily conductance as described by the reciprocal function g_{srmax for}=(-0.47+2.61._p)^-1 the evergreen oaks and g_{srmax for}=(-1.94+7.39._p)^-1 for the deciduous species. These differences between the two groups may partialy explain their geograhic distributions, and suggest general questions concerning the mechanisms which optimize water-use efficiency in Mediterranean oak species.     

The role of the tolerance–fecundity trade‐off in maintaining intraspecific seed trait variation in a widespread dimorphic herb

Coexistence of species with different seed sizes is a long‐standing issue in community ecology, and a trade‐off between fecundity and stress tolerance has been proposed to explain co‐occurrence in heterogeneous environments. Here we tested an intraspecific extension of this model: whether such trade‐off also explains seed trait variation among populations of widespread plants under stress gradients. We collected seeds from 14 populations of Plantago coronopus along the Atlantic coast in North Africa and Europe. This herb presents seed dimorphism, producing large basal seeds with a mucilaginous coat that facilitates water absorption (more stress tolerant), and small apical seeds without coats (less stress tolerant). We analysed variation among populations in number, size and mucilage production of basal and apical seeds, and searched for relationships between local environment and plant size. Populations under higher stress (higher temperature, lower precipitation, lower soil organic matter) had fewer seeds per fruit, higher predominance of basal relative to apical seeds, and larger basal seeds with thicker mucilaginous coats. These results strongly suggest a trade‐off between tolerance and fecundity at the fruit level underpins variation in seed traits among P. coronopus populations. However, seed production per plant showed the opposite pattern to seed production per fruit, and seemed related to plant size and other life‐cycle components, as an additional strategy to cope with environmental variation across the range. The tolerance–fecundity model may constitute, under stress gradients, a broader ecological framework to explain trait variation than the classical seed size–number compromise, although several fecundity levels and traits should be considered to understand the diverse strategies of widespread plants to maximise fitness in each set of local conditions.     

Water- and nutrient-use efficiency of a deciduous species, Vaccinium myrtillus, and an evergreen species, V. vitis-idaea, in a subalpine dwarf shrub heath in the southern Alps, Italy

Periodic measurements of gas‐exchange rates and determinations of foliar N and P concentrations were used for evaluating instantaneous water‐use efficiency and photosynthetic nutrient‐use efficiency in two co‐existing dwarf shrubs of different growth form (V. myrtillus, deciduous, and V. vitis‐idaea, evergreen) in a subalpine heath in the southern Alps of Italy. Those data were compared with cumulative assessments of water‐use efficiency and photosynthetic nutrient‐use efficiency obtained by measuring leaf carbon isotope discrimination in leaf tissues and by estimating nutrient resorption from senescing leaves. V. myrtillus presented higher dry‐weight based rates of net photosynthesis (A_weight) compared to V. vitis‐idaea. A_weight was positively correlated with foliar‐nutrient status and intercellular‐to‐ambient gradient in CO₂ concentrations. A_weight was, furthermore, negatively correlated with leaf specific mass. Instantaneous photosynthetic nutrient‐use efficiency did not differ between the two species but the percentages of N and P pools resorbed from senescing leaves were somewhat higher in the deciduous species. The evergreen species showed lower P concentrations in senescing leaves which indicated a higher proficiency in resorbing phosphorus compared to the deciduous species. In addition, the evergreen species achieved a higher carbon gain per unit foliar N and P, due to a longer mean residence time of both nutrients. The two species did not differ from each other with respect to both instantaneous and long‐term water‐use efficiency. This was consistent with the climatic pattern, showing no sign of water deficiency through the growing season. Current‐year V. vitis‐idaea leaves had a significantly higher Δ¹³C compared to previous‐year leaves, possibly mirroring a long term acclimation of evergreen leaves, as far as they age, to the habitat conditions in the understory where evergreen species are usually confined within mixed dwarf‐shrub communities.     

Changes in Patterns of Species Co‐occurrence across Two Tropical Cloud Forests Differing in Soil Nutrients and Air Temperature

Patterns of co‐occurrence of species are increasingly used to examine the contribution of biotic interactions to community assembly. We assessed patterns of co‐occurrence at four scales, in two types of tropical cloud forests in Hainan Island, China (tropical montane evergreen forests, TMEF and tropical dwarf forests, TDF) that varied significantly in soil nutrients and temperature. We tested if the patterns of co‐occurrence changed when we sorted species into classes by abundance and diameter at breast height (dbh). Co‐occurrence differed by forest type and with plot size, with significant species aggregation observed across larger plots in TDF and patterns of species segregation observed in smaller plots in TMEF. Analyses of differential abundance and dbh classes also showed that smaller plots in TMEF tend to have negative co‐occurrence patterns, but larger plots in TDF tend to show patterns of aggregation, suggesting competitive and facilitative interactions. This underscores the scale‐dependence of the processes contributing to community assembly. Furthermore, it is consistent with predictions of the stress gradient hypothesis that facilitation will be most important in biological systems subject to abiotic stress, while competition will be more important in less abiotically stressful habitats. Our results clearly demonstrate that these two types of tropical cloud forest exhibit different co‐occurrence patterns, and that these patterns are scale‐dependent, though independent of plant abundance and size class.     

Plant functional types broadly describe water use strategies in the Caatinga,a seasonally dry tropical forest in northeast Brazil

1. In seasonally dry tropical forests, plant functional type can be classified as deciduous low wood density, deciduous high wood density, or evergreen high wood density species. While deciduousness is often associated with drought‐avoidance and low wood density is often associated with tissue water storage, the degree to which these functional types may correspond to diverging and unique water use strategies has not been extensively tested.
2. We examined (a) tolerance to water stress, measured by predawn and mid‐day leaf water potential; (b) water use efficiency, measured via foliar δ¹³C; and (c) access to soil water, measured via stem water δ¹⁸O.
3. We found that deciduous low wood density species maintain high leaf water potential and low water use efficiency. Deciduous high wood density species have lower leaf water potential and variable water use efficiency. Both groups rely on shallow soil water. Evergreen high wood density species have low leaf water potential, higher water use efficiency, and access alternative water sources. These findings indicate that deciduous low wood density species are drought avoiders, with a specialized strategy for storing root and stem water. Deciduous high wood density species are moderately drought tolerant, and evergreen high wood density species are the most drought tolerant group.
4. Synthesis. Our results broadly support the plant functional type framework as a way to understand water use strategies, but also highlight species‐level differences.

     

Insight into the photosynthetic apparatus in evergreen and deciduous European oaks during autumn senescence using OJIP fluorescence transient analysis

Climate change is one of the major issues nowadays, and Mediterranean broadleaf species have been suggested to fill possible future gaps created by climate change in Central European forests. To provide a scientific‐based foundation for such practical strategies, it is important to obtain a general idea about differences and similarities in the physiology of Central European and Mediterranean species. In the present study, we evaluated the onset of leaf senescence of a broad spectrum of oak species under the Central European climate in a common garden experiment. Degradation of the photosynthetic apparatus of evergreen (Quercus ilex, Q. suber), semi‐evergreen (Q. × turneri, Q. × hispanica) and deciduous oaks (Q. robur, Q. cerris, Q. frainetto, Q. pubescens) was monitored as chlorophyll content and analysed chlorophyll fluorescence induction transients. In the deciduous species, a significant decline in chlorophyll content was observed during autumn/winter, with Q. pubescens showing the slowest decline. Analysis of fluorescence induction transients revealed a significant decline in quantum efficiency of the primary photochemistry and reaction centre density and later, a decrease in quantum efficiency of end acceptor reduction. Alterations in fluorescence parameters were compared to the decline in chlorophyll content, which occurred much more slowly than expected from the fluorescence data. The evergreen species showed no decline in chlorophyll content, nor different chlorophyll a fluorescence induction behaviour despite temperature falling below 0 °C. The hybrids showed intermediate behaviour between their parental evergreen and deciduous taxa.     

Fossils of the Fagaceae and their implications in systematics and biogeography

The fossil history of the Fagaceae from China and its systematic and biogeographic implications are discussed based on revisionary studies of the fossil records. No creditable macrofossil record of the Fagaceae exists in the Cretaceous deposits and all the Cretaceous microfossil reports remain equivocal and require further study. The Paleocene fossils show the appearance and diversification of the two groups corresponding to the subfamilies Fagoideae and Castaneoideae sensu Nixon. By the Eocene, all modern genera had been present. The oldest fagaceous fossils represent subfamily Fagoideae with affinities to the extant genus Trigonobalanus. The leaf fossil genus Berryophyllum, with affinities to Quercus subg. Cyclobalanopsis, has been documented by the early Eocene and might have occurred earlier than other fossils assignable to Quercus. The appearance of evergreen sclerophyllous Ouercus with entire leaves might have occurred earlier than those with toothed leaves. Deciduous, urticoid-leaved oak fossils (Quercus subg. Quercus sect. Quercus) had not appeared until the Miocene. Fossil equivalents of Trigonobalanus, Castanopsis and Lithocarpus had occurred in Europe and North America by the early Tertiary, suggesting that continuous distributions were achieved via the northern hemisphere land bridges. Three groups of evergreen sclerophyllous oaks of apparent close phylogenetic relationships occurred in the Hengduan mountains, the Mediterranean area and northwestern North America. Their fossil forms have become dominant elements of those vegetation zones since the Miocene. A shared fossil history indicates a possible biogeographic boundary formed by the ancient Mediterranean. The evidence suggests that the oaks might arrive in North America during two distinct geologic periods: evergreen sclerophyllous entire-leaved oaks appeared by the Early Tertiary, whereas thedeciduous oaks with urticoid leaves appeared in the Late Tertiary.     

Glacial refugia of temperate, Mediterranean and Ibero-North African flora in south-eastern Spain: new evidence from cave pollen at two Neanderthal man sites

Aim To locate glacial refugia of thermophilous plant species in Spain. Location Two south‐eastern Spanish Neanderthal man sites in Murcia; namely, the inland Cueva Negra del Estrecho del Río Quípar and the coastal Sima de las Palomas del Cabezo Gordo. Methods We use pollen found in cave sediments as a source of palaeobotanical and palaeoecological information. The findings are discussed with regard both to animal remains from both sites, and also to other refugia in south‐eastern Spain and elsewhere in the Iberian Peninsula. Results Both sequences show persistence of abundant mesothermophilous trees during the last glacial stage, suggesting both localities were reservoirs of phytodiversity and woodland species. At both sites, deciduous and evergreen oaks are the most abundant components, followed by a wide variety of deciduous trees and sclerophyllous shrubs, including Ibero‐North African xerothermic scrub near the coast. Conclusions Incomplete information underlies a common misapprehension that Iberian glacial refugia were confined to southernmost parts of the peninsula. A rather different picture of Quaternary refugia emerges from consideration of pollen sequences from caves (and other inputs such as macroscopic charcoal, spatial genetic structure of present‐day populations, faunal remains, and present‐day distribution of thermophilous species). This picture offers a view of numerous viable areas for woodland species in southern Spain, in addition to others in the mountain ranges, both in continental central Spain and those of northern Spain: these stretch from the Mediterranean coast of Catalonia to the westernmost extent of the Bay of Biscay.     

Fire disturbance disrupts co‐occurrence patterns of terrestrial vertebrates in Mediterranean woodlands

Aim This paper uses null model analysis to explore the pattern of species co‐occurrence of terrestrial vertebrate fauna in fire‐prone, mixed evergreen oak woodlands. Location The Erico–Quercion ilicis of the Mediterranean belt (50–800 m a.s.l.) in the Madonie mountain range, a regional park in northern Sicily (37°50′ N, 14°05′ E), Italy. Methods The stratified sampling of vertebrates in a secondary succession of recent burned areas (BA, 1–2 years old), intermediate burned areas (INT, 4–10 years old) and ancient burned areas (CNB, > 50 years old), plus forest fragments left within burned areas (FF, 1–2 years old) permitted the comparison of patterns of species co‐occurrence using a set of separate presence/absence matrices. First, the breeding avifauna derived from standardized point counts was analysed using Stone & Roberts’C‐score, and by a null model algorithm (fixed/equiprobable). Secondly, the analysis was repeated using all vertebrate species recorded in the succession. Results Sixty‐five species were recorded in the 2‐year study period in the four sample treatments. Birds were found to make up the largest component (63%) of the recorded assemblage. The BA treatment had the lowest species richness, followed in order by the small, medium and large FFs, and then by the CNBs. For both analyses (birds and total vertebrates), the C‐scores were quite small and not significantly different from those that could be expected by chance in the BA and INT burned areas; this indicates a random co‐occurrence among vertebrates of those assemblages. Contrariwise, for both analyses in the CNBs, the C‐scores were large and significantly different from the simulated indices, thereby indicating a non‐random co‐occurrence pattern (segregation) of vertebrates in the undisturbed woodlands. In addition, C‐score values for the surviving FFs show a significant aggregation of species. Main conclusions The null model analyses highlighted a new aspect of fire disturbance in Mediterranean woodland ecosystems: the disruption in patterns of co‐occurrence in the terrestrial vertebrate community. Wildfire alters community organization, inducing, for at least 10 years, a random aggregate of species. Communities re‐assemble themselves, showing the occurrence of species segregation at least 50 years after fire.     

Assessing the drought tolerance variability in Mediterranean alfalfa (Medicago sativa L.) genotypes under arid conditions

This work investigated the variability in drought tolerance under arid conditions of Mediterranean alfalfa genotypes with the overall aim to assess the main criteria that are associated with the relative tolerance and to discover the most tolerant ecotypes. For this, 16 alfalfa genotypes originating from seven countries of the Mediterranean basin were tested in an experimental station in south of Tunisia. The trial was conducted under two irrigation treatments. The first was normally irrigated by providing an amount of water corresponding to the potential evapotranspiration of the crop, and in the second with water deficit which was applied by stopping the irrigation during 8 weeks in summer. A significant decrease was observed under water deficit for biomass production, leaf stem ratio and water use efficiency. The sensitive index, in stress treatment, varies between 13.8% and 46.2% for dry yield. Results showed that some genotypes exhibited high-forage yield potential even in the presence of stress, mainly Amerist, Sardi10 and Siriver. Proline accumulation in leaves was greater in water-stressed plants, while the K⁺ osmo-regulatory role was not definite. High biomass production, accumulation of proline and constancy of K⁺ in leaves are the most important criteria for tolerant alfalfa.