Both temperature fluctuations and East Asian monsoons have driven plant diversification in the karst ecosystems from southern China

Karst ecosystems in southern China are species‐rich and have high levels of endemism, yet little is known regarding the evolutionary processes responsible for the origin and diversification of karst biodiversity. The genus Primulina (Gesneriaceae) comprises ca. 170 species endemic to southern China with high levels of ecological (edaphic) specialization, providing an exceptional model to study the plant diversification in karsts. We used molecular data from nine chloroplast and 11 nuclear regions and macroevolutionary analyses to assess the origin and cause of species diversification due to palaeoenvironmental changes and edaphic specialization in Primulina. We found that speciation was positively associated with changes in past temperatures and East Asian monsoons through the evolutionary history of Primulina. Climatic change around the mid‐Miocene triggered an early burst followed by a slowdown of diversification rate towards the present with the climate cooling. We detected different speciation rates among edaphic types, and transitions among soil types were infrequently and did not impact the overall speciation rate. Our findings suggest that both global temperature changes and East Asian monsoons have played crucial roles in floristic diversification within the karst ecosystems in southern China, such that speciation was higher when climate was warmer and wetter. This is the first study to directly demonstrate that past monsoon activity is positively correlated with speciation rate in East Asia. This case study could motivate further investigations to assess the impacts of past environmental changes on the origin and diversification of biodiversity in global karst ecosystems, most of which are under threat.     

Different combinations of morpho‐physiological traits are responsible for tolerance to drought in wild tomatoes Solanum chilense and Solanum peruvianum

Herbaceous species can modify leaf structure during the growing season in response to drought stress and water loss. Evolution can select combinations of traits in plants for efficient water use in restricted environments. We investigated plant traits that mediate adaptation and acclimation to water stress in two herbaceous drought‐tolerant species. Anatomical, morphological and physiological traits related to stems and leaves were examined under optimal watering (OW) and a long period of restricted watering (RW) in 11 accessions from three Solanaceae species (Solanum chilense, S. peruvianum and S. lycopersicum). The relationships between these traits were tested using linear regression and PCA. There were significant differences in anatomical traits between the species under both OW and RW, where leaf area correlated with stem diameter. Proline and total carbohydrates accumulated highly in S. chilense and S. peruvianum, respectively, and these osmolytes were strongly correlated with increased osmotic potential. Stomatal density varied between species but not between acclimation treatments, while stomatal rate was significantly higher in wild tomatoes. There was a strong positive relationship between stem growth rate and a group of traits together expressed as total stomatal number. Total stomata is described by integration of leaf area, stomatal density, height and internode length. It is proposed that constitutive adaptations and modifications through acclimation that mediate RW play an important role in tolerance to drought stress in herbaceous plants. The capacity for growth under drought stress was not associated with any single combination of traits in wild tomatoes, since the two species differed in relative levels of expression of various phenotypic traits.     

Stem hydraulic traits and leaf water-stress tolerance are co-ordinated with the leaf phenology of angiosperm trees in an Asian tropical dry karst forest

Background and Aims

The co-occurring of evergreen and deciduous angiosperm trees in Asian tropical dry forests on karst substrates suggests the existence of different water-use strategies among species. In this study it is hypothesized that the co-occurring evergreen and deciduous trees differ in stem hydraulic traits and leaf water relationships, and there will be correlated evolution in drought tolerance between leaves and stems.

Methods

A comparison was made of stem hydraulic conductivity, vulnerability curves, wood anatomy, leaf life span, leaf pressure–volume characteristics and photosynthetic capacity of six evergreen and six deciduous tree species co-occurring in a tropical dry karst forest in south-west China. The correlated evolution of leaf and stem traits was examined using both traditional and phylogenetic independent contrasts correlations.

Key Results

It was found that the deciduous trees had higher stem hydraulic efficiency, greater hydraulically weighted vessel diameter (D_h) and higher mass-based photosynthetic rate (A_m); while the evergreen species had greater xylem-cavitation resistance, lower leaf turgor-loss point water potential (π₀) and higher bulk modulus of elasticity. There were evolutionary correlations between leaf life span and stem hydraulic efficiency, A_m, and dry season π₀. Xylem-cavitation resistance was evolutionarily correlated with stem hydraulic efficiency, D_h, as well as dry season π₀. Both wood density and leaf density were closely correlated with leaf water-stress tolerance and A_m.

Conclusions

The results reveal the clear distinctions in stem hydraulic traits and leaf water-stress tolerance between the co-occurring evergreen and deciduous angiosperm trees in an Asian dry karst forest. A novel pattern was demonstrated linking leaf longevity with stem hydraulic efficiency and leaf water-stress tolerance. The results show the correlated evolution in drought tolerance between stems and leaves.Key words: Tropical dry forest, karst, leaf habit, hydraulic conductivity, cavitation resistance, leaf water-stress tolerance, wood density, leaf density, phylogenetic independent contrasts     

四种不同生活型植物幼苗对喀斯特生境干旱的生理生态适应性

喀斯特石漠化是我国西南喀斯特地区最严重的生态环境问题, 生境干旱是限制该地区植物生长的主要因素之一, 掌握喀斯特植被不同演替阶段不同生活型植物对干旱胁迫的适应策略有助于提高植被恢复的成功率。通过人工模拟4种干旱强度, 测定叶片水势、气体交换、叶绿素荧光、光合色素含量、渗透调节物质浓度、抗氧化酶活性以及生物量, 研究了喀斯特地区4种不同生活型植物幼苗对干旱胁迫的适应策略。这4种植物为常绿灌木火棘(Pyracantha fortuneana)、落叶灌木小果蔷薇(Rosa cymosa)、常绿乔木猴樟(Cinnamomum bodinieri)和落叶乔木圆果化香树(Platycarya longipes)。结果表明: 随着干旱程度的加深, 4种植物幼苗的叶片水势、光合能力、叶绿素含量、生物量增长、叶重比(LMR)、叶面积比(LAR)和比叶面积(SLA)逐渐下降, 而热耗散(NPQ)、类胡萝卜素与叶绿素含量比值、丙二醛含量和根重比(RMR)逐渐上升; 圆果化香树和猴樟的水分利用效率(A_n/g_s)、渗透调节物质浓度和抗氧化酶活性呈先升高后降低的趋势, 而火棘和小果蔷薇的A_n/g_s、脯氨酸含量和超氧化物歧化酶活性呈上升趋势。严重干旱下, 火棘和小果蔷薇幼苗的叶片水势和叶绿素含量下降较少, 具有较高的光合能力和生物量增长, 这主要是由于它们具有较低的SLA和LAR、较高的NPQ和A_n/g_s以及较高的渗透调节能力和抗氧化保护能力。中度干旱下, 猴樟幼苗叶片水势下降很少, LMR和LAR也较高, 脯氨酸含量和抗氧化酶活性非常高。但在严重干旱下, 其叶片水势、LMR、LAR和生物量增长大幅度下降, 最大光化学效率和光合速率也非常低, 渗透调节能力与抗氧化酶活性大幅度下降至正常水平以下。水分好的条件下, 圆果化香树幼苗具有较高的RMR以吸收充足的水分, 具有较高的LAR和叶绿素含量, 保证了生物量的大量积累。然而, 干旱胁迫致使其生物量大幅度下降, 主要是由于LMR、LAR、气体交换和叶绿素含量的大量下降以减少蒸腾面积、水分散失和对光能的吸收。研究结果表明, 火棘、小果蔷薇和猴樟幼苗主要采用耐旱策略, 其中猴樟抗严重干旱的能力较弱; 圆果化香树幼苗对干旱胁迫更为敏感, 主要采取避旱策略。     

热带喀斯特森林常绿和落叶榕树的水力特征和水分关系与抗旱策略

以热带喀斯特地区的直脉榕(Ficus orthoneura)和豆果榕(F.pisocarpa)为实验材料,研究了常绿和落叶树木枝条和叶片的解剖结构特征、光合水分特征和耐旱性的差异,目的在于探讨不同生活型榕树适应干旱生境的策略.直脉榕和豆果榕的叶片都有两层栅栏组织、游离状的海绵组织和钟乳体等旱生结构,同时叶片角质层蒸腾速率(gmin)和气孔导度(gs)相对较低.但与落叶的豆果榕相比,常绿的直脉榕的枝条木质部失去50％传导率的水势(P50)和gmin更低,表现出更保守的水分利用策略.总体上,两种榕树都表现出了对喀斯特干旱生境的良好适应,但是它们的适应策略表现出一定的差别.豆果榕通过落叶度过旱期,而直脉榕在结构和功能上比豆果榕更耐旱.抗旱策略和水分利用策略的不同导致两种榕树的生态位分异,减少了彼此间的水分竞争,有利于它们在喀斯特生境中共存.     

Correlation of drought traits and the predictability of osmotic potential at full leaf turgor in vegetation from New Zealand

Scientists do not know precisely how severe will be the impact of climate change on species. Evidence suggests that for some species, their future distributions might be jeopardized by local extinctions and drought‐induced tree mortality. Thus, we require models capable of estimating drought tolerance across many species. We can approach this goal by assessing functional traits. The trait osmotic potential at full turgor, π_O, is potentially a good drought indicator; however, few studies address its importance as a drought‐tolerance predictor and it is difficult to measure in the field with accuracy. In this work, we aim to answer the questions: which drought traits correlate with π_O?; do morpho‐anatomical traits correlate with π_O?; and which trees and shrubs are more (or less) vulnerable to drought? To achieve this aim, we assessed physiological and morpho‐anatomical traits for 14 native species from New Zealand forests. We included leaf‐ and wood‐related traits, π_O, water potential and stomatal conductance. We examined how these traits correlate with π_O and sought to generate models to predict π_O as a function of other traits. We tested 33 different models and evaluated them using Akaike's information criterion. Unfortunately, none of the morpho‐anatomical traits correlated well with π_O. Instead, water potential correlated most strongly with π_O. None of the models using only morpho‐anatomical traits produced plausible results. The model with the best predictive performance incorporated the effects of both morpho‐anatomical and physiological traits: water potential and wood saturated water content. Of the species analysed, and based on their π_O response, Lophozonia menziesii was considered the most vulnerable to drought stress, whereas Plagianthus regius was the least vulnerable. Our findings imply that it is potentially valuable to keep exploring the use of π_O as a drought indicator and that the effort required to measure some physiological traits, such as water potential, may be essential to consider plant drought responses and to predict π_O.     

Leaf anatomical structures of <Emphasis Type="Italic">Paphiopedilum</Emphasis> and <Emphasis Type="Italic">Cypripedium</Emphasis> and their adaptive significance

Paphiopedilum and Cypripedium are closely related in phylogeny, but have contrasting leaf traits and habitats. To understand the divergence in leaf traits of Paphiopedilum and Cypripedium and their adaptive significance, we analyzed the leaf anatomical structures, leaf dry mass per area (LMA), leaf lifespan (LL), leaf nitrogen concentration (N _mass), leaf phosphorus concentration (P _mass), mass-based light-saturated photosynthetic rate (A _mass), water use efficiency (WUE), photosynthetic nitrogen use efficiency (PNUE) and leaf construction cost (CC) for six species. Compared with Cypripedium, Paphiopedilum was characterized by drought tolerance derived from its leaf anatomical structures, including fleshy leaves, thick surface cuticles, huge adaxial epidermis cells, lower total stoma area, and sunken stomata. The special leaf structures of Paphiopedilum were accompanied by longer LL; higher LMA, WUE, and CC; and lower N _mass, P _mass, A _mass, and PNUE compared with Cypripedium. Leaf traits in Paphiopedilum helped it adapt to arid and nutrient-poor karst habitats. However, the leaf traits of Cypripedium reflect adaptations to an environment characterized by rich soil, abundant soil water, and significant seasonal fluctuations in temperature and precipitation. The present results contribute to our understanding of the divergent adaptation of leaf traits in slipper orchids, which is beneficial for the conservation of endangered orchids.     

干旱胁迫与复水对块根紫金牛生理特性的影响

以岩溶特有药用植物块根紫金牛为试材,研究土壤水分胁迫及复水条件下其叶片光合参数、相对含水量、质膜透性、渗透调节物质含量的变化特性。结果表明:水分胁迫下,块根紫金牛的叶片净光合速率、气孔导度和蒸腾速率均几乎接近零点,但胞间CO2浓度上升,即非气孔因素限制是光合速率下降的主要原因。水分胁迫不影响块根紫金牛单位面积的总叶绿素和类胡萝卜素含量,但干旱处理的Chl a/b和Car/Chl分别显著低于和高于对照。水分胁迫下,块根紫金牛的叶片相对含水量、相对电导率和丙二醛含量显著增大,即膜系统受到一定的伤害;块根紫金牛叶片脯氨酸含量显著降低,可溶性蛋白含量无显著变化,可溶性糖含量显著增大,但增大幅度不大,说明其在干旱胁迫下的渗透调节能力较弱。复水处理后,块根紫金牛全部指标均能恢复到对照水平,说明其对干旱胁迫较为敏感,主要采取避旱策略。     

Ecophysiological traits associated with drought in Mediterranean tree seedlings: individual responses versus interspecific trends in eleven species

Species differ regarding their drought tolerance and individuals of a given species can modify their morphology and physiology in response to drought. However, since evolutionary and ecological selective pressures differ, individual and interspecific responses to drought might not match. We determined summer survival and a number of ecophysiological variables in two factorial experiments with seedlings of eleven tree species present in Mediterranean ecosystems, grown under slowly imposed water stress and control conditions. Plants experiencing drought exhibited reduced growth, low specific leaf area, chlorophyll content, and photosynthetic rate when compared to the controls, and species-specific drought tolerance was associated with an analogous set of trait values. However, while species with high leaf area ratio and shoot-root ratio exhibited greater drought tolerance, drought induced the reversed response within species. Contrary to expectations, water use efficiency was lower in drought-tolerant species and decreased in water-stressed individuals compared to the control plants. There was a distinctive phylogenetic signal in the functional grouping of species, with oaks, pines, and other genera being clearly different from each other in their drought tolerance and in their functional responses to drought. However, all relationships between ecophysiological variables and drought tolerance were significant after accounting for phylogenetic effects, with the exception of the relationship between drought tolerance and photochemical efficiency. Our results show that drought tolerance is not achieved by a single combination of trait values, and that even though evolutionary processes and individual responses tend to render similar results in terms of functional traits associated with drought, they do not necessarily match.     

Elucidating the role of genetic drift and natural selection in cork oak differentiation regarding drought tolerance

Drought is the main selection agent in Mediterranean ecosystems and it has been suggested as an important evolutionary force responsible for population diversification in these types of environments. However, population divergence in quantitative traits can be driven by either natural selection, genetic drift or both. To investigate the roles of these forces on among-population divergence in ecophysiological traits related to drought tolerance (carbon isotope discrimination, specific leaf area, leaf size and leaf nitrogen content), we compared molecular and quantitative genetic differentiation in a common garden experiment including thirteen cork oak ( Quercus suber L.) populations across a gradient of rainfall and temperature. Population differentiation for height, specific leaf area, leaf size and nitrogen leaf content measured during a dry year far exceeded the molecular differentiation measured by six nuclear microsatellites. Populations from dry-cool sites showed the lowest nitrogen leaf content and the smallest and thickest leaves contrasting with those from humid-warm sites. These results suggest (i) these traits are subjected to divergence selection and (ii) the genetic differences among populations are partly due to climate adaptation. By contrast, the low among-population divergence found in basal diameter, annual growth and carbon isotopic discrimination (a surrogate for water use efficiency) suggests low or no divergence selection for these traits. Among-population differentiation for neutral markers was not a good predictor for differentiation regarding the quantitative traits studied here, except for leaf size. The correlation observed between the genetic differentiation for leaf size and that for molecular markers was exclusively due to the association between leaf size and the microsatellite Qp ZAG46, which suggests a possible linkage between Qp ZAG46 and genes encoding for leaf size.     

Plant ecology of tropical and subtropical karst ecosystems

Substantial areas of tropical forests, including those within nine tropical biodiversity hotspots, contain karst landscapes that have developed on soluble carbonate rocks. Here, we review how the ecology of karst forest trees is influenced by hydrological, edaphic, and topographic factors that exhibit fine spatial heterogeneity. Comparative analysis of drought tolerance traits including wood density contributes to the assessment of whether karst tree species are more drought‐tolerant compared to non‐karst trees. Although karst ecosystems are generally considered to have low phosphorus availability, foliar nitrogen‐to‐phosphorus ratios exhibit wide variation across karst regions without a clear difference from non‐karst ecosystems. According to the analyses of leaf phenology, stem water storage, and isotopic signatures from xylem sap, water use strategies of karst trees can be classified into five types: (a) soil water dependent, (b) epikarst water dependent (mainly use water stored in fine pores and gaps within the epikarst rock during the dry season), (c) groundwater dependent, (d) fog water dependent, and (e) drought‐deciduous (shed leaves during the dry season). Overall, published data suggest that only a subset of karst tree species are exclusively distributed within karst hilltops where water availability is limited. The diverse resource acquisition and utilization strategies of karst plants across edaphic habitats must be considered when developing effective strategies to conserve and restore biodiversity in karst landscapes, which are under increasing anthropogenic pressure.     

Drought effects on resource partition and conservation among leaf ontogenetic stages in epiphytic tank bromeliads

Studying the response to drought stress of keystone epiphytes such as tank bromeliads is essential to better understand their resistance capacity to future climate change. The objective was to test whether there is any variation in the carbon, water and nutrient status among different leaf ontogenetic stages in a bromeliad rosette subjected to a gradient of drought stress. We used a semi-controlled experiment consisting in a gradient of water shortage in Aechmea aquilega and Lutheria splendens. For each bromeliad and drought treatment, three leaves were collected based on their position in the rosette and several functional traits related to water and nutrient status, and carbon metabolism were measured. We found that water status traits (relative water content, leaf succulence, osmotic and midday water potentials) and carbon metabolism traits (carbon assimilation, maximum quantum yield of photosystem II, chlorophyll and starch contents) decreased with increasing drought stress, while leaf soluble sugars and carbon, nitrogen and phosphorus contents remained unchanged. The different leaf ontogenetic stages showed only marginal variations when subjected to a gradient of drought. Resources were not reallocated between different leaf ontogenetic stages but we found a reallocation of soluble sugars from leaf starch reserves to the root system. Both species were capable of metabolic and physiological adjustments in response to drought. Overall, this study advances our understanding of the resistance of bromeliads faced with increasing drought stress and paves the way for in-depth reflection on their strategies to cope with water shortage.     

Correlations among leaf structure,drought tolerance and photosynthetic capacity in saplings of Euphorbiaceae from different micro-habitats in a seasonal tropical rainforest

Aims Leaf structural and functional traits have been extensively studied to explain community assembly mechanisms, species distributions, niche differentiations, and even ecosystem services functions. However, these traits are influenced by both environment and phylogeny, showing correlations or trade-offs among them. In this study, we assessed the impacts of leaf structure on drought tolerance and photosynthetic potential, and the trade-off between drought tolerance and photosynthetic capacity, to provide an explanation for species coexistence and the maintenance of high biodiversity in tropical rainforests.
Methods We chose 18 species in the Euphobiaceae family differing in distribution patterns along topographic gradients in a 20 hm² forest dynamics monitoring plot (FDP) in Xishuangbanna. We measured leaf anatomy, leaf water relations characteristics, maximum photosynthetic rate, and dark respiration, and used two different methods—the traditional Pearson correlation and phylogenetic independent contrasts—to analyze the relationships among those traits.
Important findings We found that: 1) species showed convergence in structures and functions within specific habitat; species on ridge or slope had a stronger water loss-tolerance abilities than species in the valley. 2) Correlations among some key traits (specifically, leaf density, water potential at turgor loss point, and water potential at full turgor, etc.) were found among habitats; plants adjusted leaf structure to influence simultaneously plant water loss-tolerance abilities and photosynthetic capability, which may result in a trade-off between drought tolerance (high leaf density, leaf mass per area) and photosynthetic capability (low leaf density, leaf mass per area). 3) The phylogenetic independent contrasts must be used when analyzing correlations among the traits of geneticallyrelated species due to the weakness of traditional Pearson analysis. The ecological niche differentiation to water and light gradients as revealed by the present study provides a potential explanation for the high diversity of the seasonal tropical rainforest.     

喀斯特森林木本植物叶片功能性状对土壤特性的响应

为明确喀斯特森林植物叶片功能性状对土壤特性的响应,采用样地与样线相结合的方法调查茂兰喀斯特森林的木本植物群落,计算不同地形木本植物叶片加权平均性状值,运用单因素方差分析和冗余分析不同地形植物叶片功能性状的差异及其与土壤特性的关系。结果表明, 在生长型(常绿、落叶)和群落水平上,植物叶片功能性状在不同地形间存在显著差异(P<0.05),其中叶面积最为敏感,对生境的响应明显,常绿植物的叶厚度大于落叶植物,比叶面积则相反,而叶绿素含量差异不显著(P>0.05)。不同地形间土壤特性差异显著(P<0.05),漏斗地形土壤的田间持水量、毛管孔隙度、全氮含量、全磷含量及有机质含量较高,土壤肥力最佳,槽谷和阴坡次之,而阳坡地段土壤相对贫瘠。不同地形植物叶片功能性状与土壤特性间具有相关性,但不同地形土壤特性对叶片功能性状变异的解释率不同,影响植物叶片功能性状的主要土壤特性为有机质含量、全氮含量、全磷含量、田间持水量和土壤容重。茂兰喀斯特森林不同地形植物叶片功能性状和土壤特性的差异较大,随着土壤特性的改变,叶片功能性状的响应特征不同,这有利于林区物种共存及生物多样性维持。     

喀斯特高原峡谷优势种叶片功能性状分析

该研究对喀斯特石漠化高原峡谷地区优势种的叶片功能性状进行了分析,调查了17种植物的叶片厚度、叶面积、叶片鲜重、叶片干重、叶干物质含量、比叶面积、叶组织密度等能反映植物生存策略且易于测量的叶片功能性状,并采用逐步回归的方法探究了叶片功能性状与土壤养分之间的关系。结果表明:(1)不同优势种叶片功能性状差异明显,叶片的厚度为0.18～0.78 mm、鲜重为0.07～6.51 g、干重为0.04～3.19 g、叶面积为3.07～325.64 cm~2、叶干物质含量为318.61～573.22 mg·g~(-1)、比叶面积为60.98～236.90 cm~2·g~(-1)、叶组织密度为0.022 1～0.036 g·cm~(-3)。(2)植物通过较小的比叶面积与较大的叶干物质含量来减少水分散失、增加养分储存,以适应高温、缺水少土的环境。(3)叶片功能性状之间存在广泛的相关关系,且均达到极显著水平。叶片厚度对鲜重、干重均表现为促进效应,比叶面积与叶干物质含量及叶组织密度之间均存在极显著的抑制效应。(4)叶片功能性状随土壤养分的变化产生规律性变化。土壤全氮、全钾和有机碳对叶片功能性状产生显著影响,尤以土壤有机碳的影响更为显著。进一步分析发现,喀斯特高原峡谷地区植被恢复及重建的优势种或建群种为翅荚香槐、清香木、枇杷等;施用有机肥能够提高花椒、金银花等经济林树种的适应能力。     

热带季雨林不同小生境大戟科植物幼树的叶片结构、耐旱性和光合能力之间的相关性

植物的叶片结构和功能性状受到自身、环境和系统发育的影响。该研究选取西双版纳20 hm²热带雨林动态监测大样地内18种分布格局不同的大戟科植物, 测量了幼树叶片的解剖结构、水分关系特征、最大光合能力和暗呼吸, 主要探讨了叶片结构对植物耐旱性和光合能力的影响, 耐旱性和光合能力之间的权衡关系, 以及环境水分条件对植物功能性状相关性的影响。结果表明: 1)生境内植物表现出一定的结构和功能的趋同性, 分布在山脊和山坡的种比沟谷种具有更强的耐失水能力; 2)去除了系统发育的影响后, 一些关键性状(特别是叶片密度和膨压丧失点时的水势、饱和渗透势等)之间存在跨生境尺度上的相关关系, 植物叶片结构同时影响了植物的耐失水能力和光合能力, 植物叶片自身的结构限制导致了植物的耐旱性(高的叶片密度、比叶质量)和光合能力(低的叶片密度、比叶质量)存在反向进化关系; 3)如果研究的植物类群亲缘关系较近, 传统的Pearson相关分析不能很好地揭示其性状间的相关关系, 因而必须采用系统发育独立对照差作相关分析。大戟科植物的结构和功能在水分梯度和光梯度上的生态位分化也从功能性状的角度为热带季雨林能维持高生物多样性, 保持植物物种长期共存提供了一个可能的解释。     

Exploring the causes of variation in phenology and morphology in Mediterranean geophytes: a genus-wide study of Cyclamen

The phenology and morphology of Mediterranean plants are constrained by drought in summer and cold temperatures in winter. In this study we examine how climatic factors and phylogenetic constraints have shaped variation in the phenology and morphology of 17 species of the genus Cyclamen cultivated in uniform garden conditions. We quantify the extent to which traits differ among subgenera and thus represent conserved traits within evolutionary lineages. We also explore whether leaf, flowering and seed-release phenology are correlated among species, and thus whether variation in flowering phenology results from selection on dispersal phenology. Our results show a significant influence of subgenus membership on leaf and flowering phenology but not on morphological traits or the timing of seed release. Among-species variation in foliage height, leaf size and seed mass (but not in floral traits) is correlated with chromosome number. Leaf traits show that species with a shorter vegetative period have a higher capacity for resource acquisition. Major phenological shifts, i.e. spring vs. autumn flowering and a decoupling of leaf and flower phenology in autumnal flowering species, thus occurred prior to the diversification of species in each subgenus and not as a response to selection on dispersal timing. Leaf and flowering phenology illustrate a gradient of strategies from autumn flowering in the absence of leaves (hysteranthous species) to spring flowering with fully developed foliage (synanthous species). In the former, flowering is uncoupled from resource acquisition by simultaneous photosynthesis, indicative that hysteranthy is a response to temporal unpredictability in the onset of rain after the summer drought. Our results support the idea that whereas leaf development is controlled primarily by moisture availability and secondarily by temperature, flowering is temperature dependent, above a minimum moisture threshold. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145 , 469–484.     

Differences in xylem and leaf hydraulic traits explain differences in drought tolerance among mature Amazon rainforest trees

Considerable uncertainty surrounds the impacts of anthropogenic climate change on the composition and structure of Amazon forests. Building upon results from two large‐scale ecosystem drought experiments in the eastern Brazilian Amazon that observed increases in mortality rates among some tree species but not others, in this study we investigate the physiological traits underpinning these differential demographic responses. Xylem pressure at 50% conductivity (xylem‐P₅₀), leaf turgor loss point (TLP), cellular osmotic potential (π_o), and cellular bulk modulus of elasticity (ε), all traits mechanistically linked to drought tolerance, were measured on upper canopy branches and leaves of mature trees from selected species growing at the two drought experiment sites. Each species was placed a priori into one of four plant functional type (PFT) categories: drought‐tolerant versus drought‐intolerant based on observed mortality rates, and subdivided into early‐ versus late‐successional based on wood density. We tested the hypotheses that the measured traits would be significantly different between the four PFTs and that they would be spatially conserved across the two experimental sites. Xylem‐P₅₀, TLP, and π_o, but not ε, occurred at significantly higher water potentials for the drought‐intolerant PFT compared to the drought‐tolerant PFT; however, there were no significant differences between the early‐ and late‐successional PFTs. These results suggest that these three traits are important for determining drought tolerance, and are largely independent of wood density—a trait commonly associated with successional status. Differences in these physiological traits that occurred between the drought‐tolerant and drought‐intolerant PFTs were conserved between the two research sites, even though they had different soil types and dry‐season lengths. This more detailed understanding of how xylem and leaf hydraulic traits vary between co‐occuring drought‐tolerant and drought‐intolerant tropical tree species promises to facilitate a much‐needed improvement in the representation of plant hydraulics within terrestrial ecosystem and biosphere models, which will enhance our ability to make robust predictions of how future changes in climate will affect tropical forests.     

Limited capacity to cope with excessive light in the open and with seasonal drought in the shade in Mediterranean Ilex aquifolium populations

Climate change is expected to involve more-frequent and intense summer droughts in the Mediterranean region. This represents a threat for long-term persistence of woody species, such as European holly (Ilex aquifolium), that originated under humid climates during the Tertiary period. The capacity of this species to persist under increased water stress, both in gaps and in the understory of an oak-dominated woodland, was assessed by quantifying phenotypic plasticity in response to drought and shade. Physiological responses in plant-water relations and gas exchange were used as performance indicators under the different environments. Phenotypic plasticity of drought-stressed holly trees in response to changes in the light environment was low relative to the known response of co-occurring forest trees. Differences between morphological traits (e.g. specific leaf area and leaf: sapwood ratio in twigs) of sun- and shade-grown trees were small but significant while physiological traits were largely unresponsive to light availability. This supports the hypothesis that late-successional shade-tolerant species exhibit greater morphological than physiological plasticity. Sapling acclimation capacity through physiological mechanisms such as osmotic adjustment was insufficient to protect from summer drought. Holly mainly inhabits oceanic climates where extreme temperatures and droughts are unusual. Our results suggest that the species occupies a narrowing niche in continental Mediterranean habitats, and may lack the capacity to persist under more-severe future climate scenarios because of its low phenotypic plasticity in response to light and drought stresses.