Intraspecific variation of drought adaptation in brittlebush: leaf pubescence and timing of leaf loss vary with rainfall

Reflective leaf pubescence of the desert shrub Encelia farinosa (brittlebrush) reduces leaf temperature and plant water loss, and is considered adaptive in xeric environments. Yet, little is known about intraspecific variation in this trait. Among three populations in the northern range of E. farinosa, which span a very broad precipitation gradient, both leaf absorptance variation and differences in the timing of drought-induced leaf loss were broadly associated with climatic variability. Where mean annual rainfall was greatest, drought-induced leaf loss was earliest, but these plants also had higher population-level mean leaf absorptance values. Higher absorptance increases the relative dependence on latent heat transfer (transpirational cooling), but it also provides greater instantaneous carbon assimilation. Plants at the driest site reached lower leaf absorptance values and maintained leaves longer into the drought period. Lower leaf absorptance reduces water consumption, and extended leaf longevity may buffer against the unpredictability of growing conditions experienced in the driest site. These observations are consistent with a trade-off scenario in which plants from wetter regions might trade off water conservation for higher instantaneous carbon gain, whereas plants from drier regions reduce water consumption and extend leaf longevity to maintain photosynthetic activity in the face of unpredictable growing conditions. Received: 2 April 1997 / Accepted: 11 August 1997     

Carbon isotope discrimination differences within and between contrasting populations of<Emphasis Type="Italic"> Encelia farinosa</Emphasis> raised under common-environment conditions

Previous studies of the desert shrub Encelia farinosa have shown variation of morpholological and physiological integration that appears to match environmental differences among populations. Such findings led us to ask if there is a genetic basis for such differentiation that may be related to physiological control of intercellular CO(2) concentrations as indicated by carbon isotope discrimination (Delta) values, and if genetic variance for Delta is detectable within populations. Under common environment conditions, Delta values were compared between two populations of E. farinosa from desert regions with contrasting rainfall patterns: Superior, Ariz., a region with high annual rainfall and droughts of short duration, and Oatman, Ariz. a region with lower annual rainfall and longer drought periods. Superior plants had consistently greater mean Delta values than Oatman plants across a broad range of soil water potentials, indicating that there is a genetic basis for Delta variation between these populations. At the intrapopulation level only Oatman plants showed detectable genetic variance of Delta based on: (1) consistent individual-rank values for Delta among soil-drought stages, and (2) evidence of heritable genetic variance for Delta during one drought stage. No genetic variance in Delta was evident for the Superior population. It is hypothesized that the high spatio-temporal heterogeneity of water availability at Oatman may facilitate the maintenance of genetic variance for carbon isotope discrimination within this population. Both the inter- and intra-population level findings suggest that selection associated with rainfall and drought has resulted in genetic divergence of the physiological factors involved in Delta determination for these populations. There appears to be strong differences of water-use and carbon-gain strategies among populations, and broader functional breadth among plants in the habitat of greatest environmental heterogeneity.     

Architecture of coastal and desert Encelia farinosa (Asteraceae): consequences of plastic and heritable variation in leaf characters

The shrub Encelia farinosa (Asteraceae) exhibits geographic variation in aboveground architecture and leaf traits in parallel with environmental variation in temperature and moisture. Measurements of plants occurring across a natural gradient demonstrated that plants in desert populations produce smaller, more pubescent leaves and are more compact and branched than plants in more mesic coastal environments. This phenotypic variation is interpreted in part as adaptive genetic differentiation; small size and pubescence reduce leaf temperature and thus increase water-use efficiency but at the cost of lower photosynthetic rate, which results in slower growth and more compact growth form. We explored the basis of phenotypic variation by planting seed offspring from coastal and desert populations in common gardens in both environments. Phenotypic differences among populations persisted in both common gardens, suggesting a genetic basis for trait variation. Desert offspring outperformed coastal offspring in the desert garden, suggesting superior adaptation to hot, dry conditions. Herbivore damage was greater for all offspring in the coastal garden. Phenotypic characters also showed plastic responses; all offspring had smaller, more pubescent leaves and more compact growth form in the desert garden. Our results confirm that leaf size and pubescence are heritable characters associated with pronounced variation in plant architecture.     

Environmental Heterogeneity and Population Differentiation in Plasticity to Drought in <Emphasis Type="Italic">Convolvulus Chilensis</Emphasis> (Convolvulaceae)

Plant populations may show differentiation in phenotypic plasticity, and theory predicts that greater levels of environmental heterogeneity should select for higher magnitudes of phenotypic plasticity. We evaluated phenotypic responses to reduced soil moisture in plants of Convolvulus chilensis grown in a greenhouse from seeds collected in three natural populations that differ in environmental heterogeneity (precipitation regime). Among several morphological and ecophysiological traits evaluated, only four traits showed differentiation among populations in plasticity to soil moisture: leaf area, leaf shape, leaf area ratio (LAR), and foliar trichome density. In all of these traits plasticity to drought was greatest in plants from the population with the highest interannual variation in precipitation. We further tested the adaptive nature of these plastic responses by evaluating the relationship between phenotypic traits and total biomass, as a proxy for plant fitness, in the low water environment. Foliar trichome density appears to be the only trait that shows adaptive patterns of plasticity to drought. Plants from populations showing plasticity had higher trichome density when growing in soils with reduced moisture, and foliar trichome density was positively associated with total biomass. Co-ordinating editor: F. Stuefer     

Spatial patterns of polygenic variation in Impatiens capensis,a species with an environmentally controlled mixed mating system

Impatiens capensis displays a mixed mating system in which individual out-crossing rate is expected to increase with light and resource availability. We investigated the amount and spatial distribution of polygenic variation for 15 morphological traits within and among six natural populations of I. capensis growing in three distinct light habitats (shaded, mixed, full sun). We grew individuals from each population in uniform greenhouse conditions and detected significant genetic variation among families within populations for all the quantitative traits examined. However, only the features related to the vegetative characteristics of seedlings and sexually mature plants show also differentiation at the population level. Surprisingly, even though light availability is likely to be the most important factor affecting the mating system of I. capensis, we find that: (1) trait means of individuals from similar light environments are not more similar than those from different light environments; (2) partitioning of polygenic variance within and among families differs both among populations from the same light habitat and among characters within each population. If natural selection is maintaining such variation, it must operate primarily through heterogeneous selection pressure within, rather than between, populations.     

Population variation in plant traits associated with ant attraction and herbivory in Chamaecrista fasciculata (Fabaceae)

The benefits of ant–plant–herbivore interactions for the plant depend on the abundance of ants and herbivores and the selective pressures these arthropods exert. In plants bearing extrafloral nectaries (EFN), different mean trait values may be selected for by different populations in response to local herbivore pressure, ultimately resulting in the evolution of differences in plant traits that attract ants as defensive agents against herbivory. To determine if variation in traits that mediate ant–plant interactions reflect herbivore selective pressures, we quantified intra- and inter-population variation in plant traits for eight populations of the EFN-bearing annual Chamaecrista fasciculata (Michx.) (Fabaceae). Censuses in rural and urban areas of Missouri and Illinois (USA) showed population differences in ant attendance and herbivore pressure. Seeds were collected from each population, and plants were grown in a common greenhouse environment to measure sugar production, nectar volume and composition, EFN size and time of emergence, leaf pubescence, and leaf quality throughout plant development. Populations varied mainly in terms of nectary size, sugar production, and nectar volume, but to a lesser degree in leaf pubescence. Populations of C. fasciculata within urban areas (low in insect abundance) had small nectaries and the lowest nectar production. There was a positive correlation across populations between herbivore density and leaf damage by those herbivores on the one hand and sugar production and nectar volume on the other. These results, in conjunction with lack of evidence for maternally based environmental effects, suggest that population differences in herbivore damage have promoted differential evolution of EFN-related traits among populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

浙江楠种群表型变异

以探讨浙江楠(Phoebe chekiangensis)种群间和种群内的表型变异程度和变异规律为目的, 对其分布区9个种群的10个表型性状进行了研究。采用巢式方差分析、变异系数、多重比较和相关分析等多种分析方法, 对种群间和种群内的表型多样性及其与地理生态因子的相关性进行了讨论。研究结果表明: 浙江楠叶片、种子等表型性状在种群间和种群内均存在着丰富的变异, 10个性状在种群间、种群内的差异均达极显著水平; 种群内的变异(21.74%)大于种群间的变异(18.45%), 表型性状的平均分化系数为41.43%, 种群内变异是浙江楠表型变异的主要变异来源。各性状平均变异系数为12.78%, 变化幅度为6.50%-19.38%。种群间叶片的平均变异系数(16.99%)高于种群间种子的平均变异系数(8.58%), 表明种子的稳定性高于叶片。叶宽、叶面积与叶片其他性状间多呈显著或极显著相关, 种子千粒重与种子宽、种子体积显著正相关; 而种子和叶片的各性状间多无显著相关性。种子宽、种子体积和种子千粒重随着海拔的增加出现减小的趋势, 种子体积受纬度控制, 随着纬度的增大表现出增大的趋势。     

栓皮栎叶片解剖结构种群变异及其对环境因子的适应

【目的】厘清天然广布树种栓皮栎叶片解剖结构的种群变异及其与环境因子的关系。【方法】以中国28个天然分布栓皮栎种群叶片为研究对象,用常规石蜡切片结合光学显微技术对栓皮栎叶片解剖性状进行研究,用巢式方差分析、Pearson相关性分析等方法对栓皮栎叶片解剖结构的种群变异与环境因子关系进行分析。【结果】(1)栓皮栎叶片解剖性状在28个种群间存在显著差异,平均变异系数为7.84%～15.16%,且同一叶片解剖性状在不同种群间的变异范围不同;(2)9个解剖性状在种群内及种群间均存在极显著差异,平均表型分化系数为37.44%,解剖性状变异主要来自种群内;(3)叶片厚度、上表皮厚度、栅栏组织厚度、海绵组织厚度与纬度呈显著正相关关系,年均气温及年降水量对上表皮厚度和栅栏组织厚度有显著影响。【结论】栓皮栎具有丰富的遗传多样性,为适应低温与干旱,栓皮栎叶片整体呈增厚趋势。研究结果为了解栓皮栎环境适应策略提供理论依据。     

Quantitative variation within and among populations of Arabis serrata Fr. & Sav. (Brassicaceae)

Phenotypic and genetic variation within and among eight populations of Arabis serrata are documented in this study. This species shows great morphological variation throughout its geographical distribution in Japan. Plants are located in habitats with different types of soils and degree of disturbance. Half-sibs progenies from eight populations were collected and cultivated in a garden experiment. Nine morphological traits representing size and shape of rosette leaves were recorded. Univariate analyses of measured traits showed that phenotypic means differed among populations for all characters. Leaves of plants from disturbed habitats had the longest petioles (lanceolate) and plants from limestone habitats showed the most roundness in leaf shape (ovate). The northernmost populations always revealed the smallest leaves. Multivariate principal component analyses also showed that leaf shape and size varied among populations. The first three principal components explained 98.5% of the variation. Coefficients of variation had a very wide range and differed from one population to another. Some traits (e.g. leaf width/leaf length ratio) were consistently less variable while others (e.g. leaf area and petiole length) were more plastic. All traits had significant genetic variance in all populations. Intra-class correlation coefficients differed for most of the traits and each population presented a different range of values. Most of the leaf traits were intercorrelated in all the populations studied, although some populations were integrated more tightly for some traits. Populations of A. serrata are differentiated in phenotypic means but they display a mosaic of traits with slight morphological differences in each locality (i.e. a quantitative genetic variation). Some traits can be correlated to the habitats that they occupy but for some of them it is difficult to assign an actual adaptive value.     

Variation in tolerance to drought among Scandinavian populations of <Emphasis Type="Italic">Arabidopsis lyrata</Emphasis>

The ability to cope with water limitation influences plant distributions, and several plant traits have been interpreted as adaptations to drought stress. In Scandinavia, the perennial herb Arabidopsis lyrata occurs in open habitats that differ widely in climate and water availability in summer, suggesting differential selection on drought-related traits. We conducted two greenhouse experiments to examine differentiation in drought response traits among six Scandinavian populations, and to determine whether leaf trichomes confer protection against drought. We quantified tolerance to drought as fitness (survival and biomass of survivors) when exposed to drought relative to fitness under non-drought conditions. Two Swedish populations from shores along the Bothnian Bay had higher tolerance to drought than four riverbed populations from Norway. Under conditions of drought, the shore populations experienced less leaf damage compared to the riverbed populations, and their survival and biomass were less reduced relative to non-drought conditions. Across populations, tolerance to drought was positively related to leaf mass per area and negatively related to flowering propensity and proportion roots, but not related to plant size at the initiation of the drought treatment. In populations polymorphic for trichome production, trichome-producing plants were more tolerant to drought than glabrous plants. The results suggest that both leaf morphology and life-history traits contribute to differential drought response in natural populations of A. lyrata, and that this system offers excellent opportunities for examining the adaptive value and genetic basis of drought-related traits.     

Variation in and adaptive plasticity of flower size and drought-coping traits

Although phenotypic plasticity of morphological and physiological traits in response to drought could be adaptive, there have been relatively few tests of plasticity variation or of adaptive plasticity in drought-coping traits across populations with different moisture availabilities. We measured floral size, vegetative size, and physiological traits in four field populations of Leptosiphon androsaceus (Polemoniaceae) that were distributed across a rainfall gradient in California, USA. Measurements were made over 5 years that varied in precipitation. We also conducted a growth chamber experiment in which half-sibs from three populations were divided equally among a well-watered and a drought treatment. We tested for selection on traits in each of the watering treatments, and evaluated whether traits exhibited plasticity. In the field, plant traits exhibited substantial variation across populations and years. Flower size, leaf size, and water-use efficiency (WUE) were generally higher for populations that received greater average rainfall. However, in dry years, we observed a decrease in flower and leaf size, but an increase in WUE across the populations. In the growth chamber experiment, leaf and physiological traits exhibited plasticity, with smaller leaves and higher WUE found in the drought, as compared to the well-watered treatment. Only specific leaf area exhibited differentiation in plasticity among populations. Although there was no observed plasticity in floral size, selection favored smaller flowers in the drought treatment and larger flowers in the well-watered treatment. Our results suggest that moisture availability has led to trait variation in L. androsaceus via a combination of selection and phenotypic plasticity.     

Patterns of intraspecific trait variation along an aridity gradient suggest both drought escape and drought tolerance strategies in an invasive herb

Background and AimsIn water-limited landscapes, some plants build structures that enable them to survive with minimal water (drought resistance). Instead of making structures that allow survival through times of water limitation, annual plants may invoke a drought escape strategy where they complete growth and reproduction when water is available. Drought escape and resistance each require a unique combination of traits and therefore plants are likely to have a suite of trait values that are consistent with a single drought response strategy. In environments where conditions are variable, plants may additionally evolve phenotypically plastic trait responses to water availability. Invasive annual species commonly occur in arid and semi-arid environments and many will be subject to reduced water availability associated with climate change. Assessing intraspecific trait variation across environmental gradients is a valuable tool for understanding how invasive plants establish and persist in arid environments.MethodsIn this study, we used a common garden experiment with two levels of water availability to determine how traits related to carbon assimilation, water use, biomass allocation and flowering phenology vary in California wild radish populations across an aridity gradient.Key ResultsWe found that populations from arid environments have rapid flowering and increased allocation to root biomass, traits associated with both drought escape and tolerance. Early flowering was associated with higher leaf nitrogen concentration and lower leaf mass per area, traits associated with high resource acquisition. While trait values varied across low- and high-water treatments, these shifts were consistent across populations, indicating no differential plasticity across the aridity gradient.ConclusionsWhile previous studies have suggested that drought escape and drought resistance are mutually exclusive drought response strategies, our findings suggest that invasive annuals may employ both strategies to succeed in novel semi-arid environments. As many regions are expected to become more arid in the future, investigations of intraspecific trait variation within low water environments help to inform our understanding of potential evolutionary responses to increased aridity in invasive species.     

Plasticity of inflorescence traits in Lobelia siphilitica (Lobeliaceae) in response to soil water availability

Many workers have demonstrated a genetic basis for variation in inflorescence traits, but this variation can also have an environmental component. Because flowering can incur significant water costs, I estimated plasticity of inflorescence traits of three populations of Lobelia siphilitica in response to drought. I manipulated soil water availability in the greenhouse and measured seven inflorescence traits. Under drought conditions, plants from one population flowered later and produced fewer flowers with shorter corollas and narrower landing pads. In contrast, the height of the flowering stalk decreased in response to drought in all three populations. Consequently, pollinator-mediated natural selection on these plastic traits may depend on soil water availability. Plastic responses differed between genotypes only for the height of the flowering stalk and the length of the corolla tube and only in one or two populations. This suggests that genotype × environment interactions would not limit the evolution of inflorescence traits in L. siphilitica. The strength and sign of phenotypic correlations among inflorescence traits did not respond plastically to drought, suggesting that indirect selection on inflorescence traits of L. siphilitica will not vary strongly with water availability. My results suggest that plasticity of inflorescence traits may influence their evolution, but the effects are population- and trait-specific.     

Exploring the impact of neutral evolution on intrapopulation genetic differentiation in functional traits in a long-lived plant

Most plant species, particularly long-lived plants, harbor a large amount of genetic variation within populations. A central issue in evolutionary ecology is to explore levels of genetic variation and understand the mechanisms that influence them. In this study, our goals were to examine the impact of neutral evolutionary processes on the genetic variance and functional diversity within three populations of a long-lived plant (Quercus suber L.). For this purpose, we genotyped the progeny of 45 open-pollinated mother trees from three populations originating from Spain, Portugal, and Morocco using six microsatellite markers. Seedlings were planted in a common garden trial and were phenotypically characterized by seven leaf functional traits. Molecular analyses revealed weak genetic differences between Iberian and Moroccan populations. Nevertheless, high genetic differentiation was observed among maternal families within populations. Differentiation between particular maternal families from the same population reached values of 29.2 %, which far exceeds the values reported between the most genetically distant populations for this species (11.7 %). Maternal families differed also in phenology, leaf size, and shape traits. In the Moroccan population, there were correlations among matrices of distances for molecular markers, leaf shape traits (e.g., leaf circularity index), and phenology, indicating that maternal families with contrasting phenologies were genetically and functionally distinct. This, together with the moderate heritability for phenology in Moroccan population, suggests that besides selective forces, neutral evolutionary processes have promoted intrapopulation genetic divergence and contribute to maintain high levels of genetic variation within this population. Overall, our results reinforce the importance of intrapopulation studies in long-lived plants under an evolutionary context.     

Pubescence and leaf spectral characteristics in a desert shrub,Encelia farinosa

Summary The effects of leaf hairs (pubescence) on leaf spectral characteristics were measured for the drought-deciduous desert shrub Encelia farinosa. Leaf absorptance to solar radiation is diminished by the presence of pubescence. The pubescence appears to be reflective only after the hairs have dried out. There are seasonal changes in leaf absorptance; leaves produced at the beginning of a growing season have high absorptances, whereas leaves produced during the growing season are more pubescent and have lower absorptances. The decrease in leaf absorptance is the result of an increase in pubescence density and thickness. Between 400 and 700 nm (visible wavelengths), pubescence serves as a blanket reflector. However, over the entire solar spectrum (400–3000 nm), the pubescence preferentially reflects near infrared radiation (700–3000 nm) over photosynthetically useful solar radiation (400–700 nm). Leaf absorptance to solar radiation (400–3000 nm) varies between 46 and 16%, depending on pubescence; whereas leaf absorptance to photosynthetically useful radiation (400–700 nm) may vary from 81 to 29%.C.I.W.-D.P.B. Publication No. 612     

自然干旱梯度下的酸枣表型变异

表型变异是植物应对环境变化的一种策略。酸枣植物从中国东部沿海到内陆腹地均有分布,其表型性状的变异可能解释其对自然干旱梯度的适应机制。为验证这一假说,以烟台、石家庄、银川、吐鲁番4个自然干旱梯度生境中生长的酸枣三年生植株的12个表型性状为调查研究对象,采用变异系数和巢式方差分析对酸枣的表型变异进行分析。结果表明:(1)从烟台到银川,叶面积、叶长、叶周长和叶柄长总体呈减小的趋势,而比叶面积呈增大的趋势;(2)随着干旱程度的增强,二次枝的长度、二次枝的基部粗、二次枝的枣吊数、茎比密度和茎水分含量均呈减小趋势,并且种子重和种子短轴长也均呈减小的趋势;(3)对沿干旱梯度分布的4个酸枣种群而言,叶性状的平均变异系数(33.70%)枝性状的平均变异系数(32.41%)种子性状的平均变异系数(9.07%),并且酸枣性状间存在很强的协变。结果表明酸枣的地上部分形态性状沿干旱梯度表现出很强的变异,推测在未来的气候变化下,酸枣将通过这种表型变异的有效组合来适应环境变化。     

Drought affects the coordination of belowground and aboveground resource‐related traits in Solidago canadensis in China

Quantifying patterns of variation and coordination of plant functional traits can help to understand the mechanisms underlying both invasiveness and adaptation of plants. Little is known about the coordinated variations of performance and functional traits of different organs in invasive plants, especially in response to their adaptation to environmental stressors. To identify the responses of the invasive species Solidago canadensis to drought, 180 individuals were randomly collected from 15 populations and 212 ramets were replanted in a greenhouse to investigate both the response and coordination between root and leaf functional traits. Drought significantly decreased plant growth and most of the root and leaf functional traits, that is, root length, surface area, volume and leaf size, number, and mass fraction, except for the root length ratio and root mass fraction. Phenotypic plasticity was higher in root traits than in leaf traits in response to drought, and populations did not differ significantly. The plasticity of most root functional traits, that is, root length (RL), root surface area (RSA), root volume (RV), and root mass fraction (RMF), were significantly positively correlated with biomass between control and drought. However, the opposite was found for leaf functional traits, that is, specific leaf area (SLA), leaf area ratio (LAR), and leaf mass fraction (LMF). Drought enhanced the relationship between root and leaf, that is, 26 pairwise root–leaf traits were significantly correlated under drought, while only 15 pairwise root–leaf traits were significantly correlated under control conditions. Significant correlations were found between biomass and all measured functional traits except for leaf size. RV, root length ratio, RMF, total area of leaves, and LMF responded differently to water availability. These responses enable S. canadensis to cope with drought conditions and may help to explain the reason of the vast ecological amplitude of this species.     

青藏高原特有属——固沙草属表型变异及其对环境因子的响应

该研究通过测量固沙草属(Orinus)3个物种(青海固沙草、固沙草和居间固沙草)的40个天然居群145个个体的22个表型性状,依据VIF膨胀系数筛选得到年平均降雨(MAP)等9个气候因子和海拔等因子,并采用冗余分析(db-RDA)考察不同物种、不同居群植物表型多样性与多元地理气候环境的关系,探讨该属植物表型多样性驱动机理。结果显示:(1)基于22个植物表型性状,固沙草属天然居群在组内最小方差(Ward)55时可归为2类,居群间表型性状不存在显著的空间自相关性(P 0.05),植物表型多样性高(Shannon-Wiener index,H:1.045～2.734),17个定量的表型性状变异系数(CV)平均值47.84%,而居群间小穗颜色多样性最低(H:0.170),证明固沙草属植物表型性状具有独特的环境响应模式。(2)巢式方差分析显示,固沙草属植物表型在居群间和居群内均存在丰富的变异(F 10,P0.01),旗叶、颖片和外稃分别是主成分分析中表型总变异的重要组成部分,表型分化系数居群间变异(71.10%)大于居群内(28.90%),固沙草属植物表型变异来源于居群间。(3)冗余分析结果表明,海拔(ALT)和年平均风速(Wind)是决定青海固沙草(O.kokonoricus)和固沙草(O.thoroldii)居群间表型差异性的主要影响因子;极端气候条件[最冷季平均温度(Bio9)和最干季降雨量(Bio14)]对固沙草居群间表型相似性起促进作用,相反干燥度指数(AI)与水气压(Vapr)对其差异性起促进作用;居间固沙草(O.intermedius)居群间表型差异受年均降雨(MAP)、潜在蒸散(PET)和水气压(Vapr)影响。研究认为,固沙草属植物天然居群之间表型变异反映出明显的地理气候变化趋势,这些表型变异是高山植物环境适应性的重要标志。     

Past climatic fluctuations are associated with morphological differentiation in the cloud forest endemic tree <Emphasis Type="Italic">Ocotea psychotrioides</Emphasis> (Lauraceae)

Pleistocene glacial periods have had a major influence on the geographical patterns of genetic structure of species in tropical montane regions. However, their effect on morphological differentiation among populations of cloud forest plants remains virtually unexplored. Here, we address this question by testing whether geographical patterns of morphological variation in Ocotea psychotrioides can be explained by the intensity of climate change occurring during 130,000 years. For this, we measured vegetative and reproductive traits for 96 individuals from 36 localities registered across the species’ distribution range. Species distribution models and multivariate statistics were used to investigate geographical patterns of morphological variation and test their association with current and past climatic conditions. Leaf size and pubescence in O. psychotrioides showed a latitudinal pattern of clinal variation that does not fit the geographical gradient of increasing leaf size towards lower latitudes observed globally among plants. Instead, the observed clinal variation conforms to a pattern of increasing leaf size towards higher latitudes. However, our analyses showed weak to non-significant association between morphology and current climate. Interestingly, our analyses showed that predicted shifts in the distribution range of O. psychotrioides during the last 130,000 years were accompanied by significant changes in climatic conditions, particularly temperature seasonality and precipitation. Accordingly, climatic instability showed a better fit to the observed patterns of leaf size and pubescence variation than current climate conditions. These results suggest that climatic instability during the Pleistocene glacial periods might play a key role in promoting morphological differentiation among populations of cloud forest plants.     

The temporal response to drought in a Mediterranean evergreen tree: comparing a regional precipitation gradient and a throughfall exclusion experiment

Like many midlatitude ecosystems, Mediterranean forests will suffer longer and more intense droughts with the ongoing climate change. The responses to drought in long‐lived trees differ depending on the time scale considered, and short‐term responses are currently better understood than longer term acclimation. We assessed the temporal changes in trees facing a chronic reduction in water availability by comparing leaf‐scale physiological traits, branch‐scale hydraulic traits, and stand‐scale biomass partitioning in the evergreen Quercus ilex across a regional precipitation gradient (long‐term changes) and in a partial throughfall exclusion experiment (TEE, medium term changes). At the leaf scale, gas exchange, mass per unit area and nitrogen concentration showed homeostatic responses to drought as they did not change among the sites of the precipitation gradient or in the experimental treatments of the TEE. A similar homeostatic response was observed for the xylem vulnerability to cavitation at the branch scale. In contrast, the ratio of leaf area over sapwood area (LA/SA) in young branches exhibited a transient response to drought because it decreased in response to the TEE the first 4 years of treatment, but did not change among the sites of the gradient. At the stand scale, leaf area index (LAI) decreased, and the ratios of stem SA to LAI and of fine root area to LAI both increased in trees subjected to throughfall exclusion and from the wettest to the driest site of the gradient. Taken together, these results suggest that acclimation to chronic drought in long‐lived Q. ilex is mediated by changes in hydraulic allometry that shift progressively from low (branch) to high (stand) organizational levels, and act to maintain the leaf water potential within the range of xylem hydraulic function and leaf photosynthetic assimilation.