高山植物叶片δ 13C的海拔响应及其机理

植物13C的分辨研究已成为植物生态学和全球碳循环研究的核心问题之一.植物13C的分辨是环境和生物因子共同作用的综合结果,海拔梯度变化不仅可以造成植物生存环境的变化,而且还可以造成植物形态和生理特征的变化,因此,高山植物13C分辨随海拔的变化为深入揭示植物13C分辨的环境和生物因子的作用机理提供了非常理想的研究条件.在简单介绍植物13C分辨基本理论的基础上,对目前国际上高山植物13C分辨的海拔响应研究进行了述评.重点介绍了随海拔变化的大气13C组成、温度、气压、水分等环境因子和植物叶片的气孔导度、羧化效率、氮含量和叶肉细胞导度等生物因子对高山C3植物13C分辨的影响,指出高山植物13C分辨的海拔响应机理仍存在一些不确定性,为国内相关研究的开展提供了一定参考.     

巴郎山异型柳叶片功能性状及性状间关系对海拔的响应

在卧龙自然保护区,按海拔梯度选择了4个异型柳分布地点(2350 m、2700 m、3150 m和3530 m),对各研究地点异型柳进行了叶片光合、CO2扩散导度(气孔导度(gs)和叶肉细胞导度(gm))、δ13C、氮素以及比叶面积(SLA)等参数的测量,以期揭示该植物叶片功能性状及功能性状间关系的海拔响应情况.结果表明:随着海拔的升高,大气温度和压强的降低,异型柳的叶片单位面积氮含量(Narea)、最大羧化速率(Vcmax)和最大净光合速率(Amax)均随之增加,这可能是该落叶灌木对于生长季节缩短的一种响应;同时,植物的光合氮利用效率(PNUE)和SLA却均随海拔降低,原因可能在于随着海拔的升高,植物将越来越多的氮素用于细胞壁等非光合组织的构建,这是高海拔植物对于外界恶劣环境的一种适应;最后,扩散导度和羧化能力是植物叶片δ13C的主要影响因子,而羧化能力较扩散导度对于异型柳叶片δ13C的作用更大些,进而导致该值呈现随海拔升高的趋势.氮素在光合与非光合系统间的分配是巴郎山异型柳适应不同海拔生境的关键.     

基于FvCB模型的叶片光合生理对环境因子的响应研究进展

为提高叶片光合速率并更好地理解叶片光合生理对环境因子变化的响应机制,FvCB模型(C_3植物光合生化模型)常用于分析不同环境条件下CO_2响应曲线并预测叶片活体内光合系统的内在变化状况。系统介绍了FvCB模型的建立、发展过程和拟合方法等基本理论,综述了该模型在叶片光合生理对光、CO_2、水、温度和N营养等环境因子变化的响应机制中的应用研究。为进一步完善FvCB模型并更好地理解叶片活体内光合系统对环境因子变化的响应机制,未来拟加强以下研究:1)羧化速率与光合电子传递速率之间的联系;2)叶肉导度的具体组分及其对FvCB模型参数估计的影响;3)叶片气孔导度和叶肉导度对环境因子变化的调控机制。     

3种高山植物紫外吸收物质含量随海拔与季节的变化特征

在植物生长季,对生长于不同海拔高度的3种高山植物矮嵩草(Kobresia humilis)、珠芽蓼(Polygonum viviparum)和平车前(Plantago depressa)的紫外吸收物质含量的季节和海拔变化特征进行了比较研究,以揭示高山植物对极端环境适应的生理生态学机制.结果显示:(1)在整个生长季,3种高山植物叶片紫外吸收物质含量在同时期同海拔条件下表现为平车前最低,珠芽蓼最高.随着海拔升高,紫外吸收物质含量在3种植物中均呈现增加趋势,并与海拔高度存在正相关关系.(2)3种高山植物紫外吸收物质含量随季节均呈现单峰变化趋势;矮嵩草和珠芽蓼的紫外吸收物质含量在草盛初期的6月份达到最大值,而后逐渐降低,并且与紫外辐射强度的季节动态存在一定正相关性;平车前紫外吸收物质含量在草盛期的7月份达到最大值,而后含量逐渐降低,其紫外吸收物质含量的季节变化与紫外辐射强度的季节变化呈较低的负相关,而与植物发育时期可能有关.可见,3种高山植物叶片紫外吸收物质含量随海拔与季节变化特征存在差异,这可能与它们的遗传特性和适应高原强紫外辐射环境胁迫策略有关.     

太白山栎属树种气孔特征沿海拔梯度的变化规律

气孔是植物与外界环境进行水分和气体交换的主要通道,调节植物碳同化和水分散失的平衡关系,在一定程度上反映植物对外界环境变化的适应。沿太白山北坡1100—2300 m海拔,测定4种栎属树种的气孔性状,分析气孔性状沿海拔的变化规律和其对环境因子的响应。结果表明:(1)气孔密度与气孔长度间的负相关在4个树种间均显著存在(P0.05);除栓皮栎(Quercus variabilis)外,气孔密度与潜在气孔导度指数的正相关关系均达显著水平;而气孔宽度与气孔长度之间只在栓皮栎和锐齿栎(Q. aliena var. acuteserrata)达到显著水平。(2)栓皮栎和槲栎(Q. aliena)的气孔长度和宽度随海拔升高而下降,气孔密度、潜在气孔导度指数增加,辽东栎(Q. wutaishansea)变化形式则相反;锐齿栎气孔宽度减小,其余性状沿海拔呈单峰变化,在约1600 m处气孔长度达到最小值,气孔密度和潜在气孔导度指数达到最大值。(3)与土壤因子相比,气孔性状主要受气候因素的影响。潜在气孔导度指数与大气温度、空气湿度成极显著正相关(P0.01),与降水量显著负相关(P0.05)。其中,空气相对湿度是影响潜在气孔导度指数的主要因素,能够解释气孔变异的22.9%。本研究结果对于深入认识秦岭太白山地区栎属树种对环境变化的响应和适应提供理论证据。     

太白山栎属树种叶片生态化学计量特征沿海拔梯度的变化规律

植物叶片碳（C）、氮（N）、磷（P）化学计量学能够反映植物对环境的适应性以及环境变化对植物的影响,是生态化学计量学的热点之一。研究亲缘关系相近物种对环境变化的适应差异对于深入了解植物的化学计量策略具有重要意义,而目前对于亲缘关系相近物种沿海拔梯度各如何变化未获得一致性的结论。因此,本研究在秦岭太白山海拔约1100-2200 m范围内,对槲栎（Quercus aliena）、栓皮栎（Q.variabilis）、锐齿栎（Q.aliena var.acuteserrata）、辽东栎（Q.wutaishansea）这4种栎属树种的叶片C、N、P含量进行测定与分析,考察叶片化学计量特征随海拔的变化趋势,同时量化气候、土壤和地形3种影响因素对其变异规律的解释程度。结果表明：（1）总体来看,4树种叶片C含量随海拔升高先上升后降低,叶片N含量和N：P则表现出随海拔升高而降低的趋势,而叶片C：N随海拔升高而升高。（2）不同树种随海拔的变化趋势不同：槲栎与锐齿栎具有相似性,叶片N、P含量都随海拔升高显著降低,C：N都随海拔升高显著升高;栓皮栎的叶片N含量和C：N与前两者呈现相反趋势;辽东栎叶片C含量随海拔上升而下降,与栓皮栎相同,但其叶片P含量和N：P分别呈现先升高后降低、先降低后升高的曲线变化趋势。（3）叶片不同化学计量特征值受到不同因子的影响。其中,叶片N含量和C：N主要受气候因子影响（解释度为39.91%和36.59%）;叶片C含量主要受土壤因子影响（解释度为25.22%）;叶片P含量、N：P和C：P则主要受到土壤因子和坡度影响（解释度有23.70%-39.83%）,且这两个因子的交互效应影响较大（交互效应解释度有16.24%-24.72%）。本研究结果说明：（1）亲缘关系较近的物种在应对环境变化时,也会有不同的变化格局及对应的养分策略,而且这能在一定程度上解释它们的地带性分布规律;（2）地形因子会与土壤因子共同影响植物的化学计量特征,在研究山地森林生态系统时,坡度也是需要考虑的重要影响因子。     

青藏高原东部典型高山植物叶片δ13C的季节变化

通过对青藏高原高寒草甸生态系统28种高山植物叶片不同月份稳定碳同位素组成的测定,研究植物δ 13C值在不同季节变化及其与环境之间的关系,试图找出影响δ 13C值变化的关键环境因子.结果表明植物δ13C值在不同月份间有显著性差异(P<0.01),生长初期(6月)δ13C值明显高于生长末期(8月).植物的δ13C值变化主要是由于温度和降水引起的,随温度和降雨量降低而偏重.另外,不同生长期植物叶片的成熟度可能对植物δ13C的变化有一定的贡献.不同种植物稳定性碳同位素值变化差别很大,反应了不同植物对环境变化的不同响应.     

羊草叶片气孔导度对环境因子的响应模拟

准确定量描述植物气孔对环境的响应是了解植物光合作用机理、预测植物生产力及其大气－植被－土壤系统中水分和热量交换的关键。利用松嫩平原盐碱化草地羊草光合生理特征的野外观测数据,分析了羊草叶片气孔导度对环境因子的反应,结果表明：羊草叶片气孔导度对环境因子变化敏感,尤其对瞬时光合有效辐射（ＰＡＲ）、叶片与空气间的水汽压亏损（ＶＰＤ）和空气温度（Ｔａ）反应十分明显。依据野外实测资料对国际上两类代表性气孔导度     

不同海拔华北落叶松细根形态特征

山地海拔变化包含多个环境因子的梯度效应,而细根作为植物重要的功能器官,对环境因子变化较为敏感。因此,了解植物细根形态特征对海拔变化的响应对于认识气候变化下的植物地下过程具有重要意义。本研究以山西省庞泉沟国家自然保护区内分布于1800~2700 m海拔上的华北落叶松(Larix principis-rupprechtii)细根为研究对象,采用根序分级法对不同根序细根形态特征(直径、比根长、比表面积和组织密度)进行了分析,结果表明:(1)同级根序细根直径随海拔升高而增粗,组织密度随海拔升高而减小,比根长和比表面积则随海拔升高先增加后减小;(2)不同海拔处细根形态均表现随序级增加,直径和组织密度变大,而比根长和比表面积减小;(3)分析表明,海拔、根序变化均显著影响细根形态特征(P0.05),但海拔和根序的交互作用只对细根直径和组织密度变化有显著影响(P0.05);其中1~3级根直径和组织密度均与海拔变化显著相关(P0.05),而4、5级根序各形态特征与海拔之间的相关性均不显著。本研究结果可为进一步认识植物细根对未来气候变化下的响应机制提供重要参考。     

桃树冠层蒸腾动态的数学模拟

将气孔导度公式、Penman—Monteith公式和土壤水分限制模型相结合,可以模拟出不同环境因子对植物蒸腾进程的影响。通过对盆栽桃树（Prunus persica var．nectadna Maxim．）数值模拟发现：影响桃树蒸腾速率的主要气象因子是太阳辐射、大气温度和湿度。植物通过气孔导度的改变来响应气象因子的变化,蒸腾的日变化主要是由气象因子的日变化引起的。土壤的水分状况也对气孔导度有显著的影响,进而影响植物的蒸腾大小。通过数值模拟还发现植物的蒸腾量并不总是随叶面积的增大而增大,对于桃树而言叶面积指数为4左右时日蒸腾量达到最大值。通过对气孔导度和蒸腾速率的模拟值和实测值进行检验发现,两者基本吻合,说明利用数学模拟的方法可以求出不同环境条件和不同叶面积桃树冠层的蒸腾速率。     

Contrasting altitudinal variation of alpine plant communities along the Swedish mountains

Changes in abiotic factors along altitudinal and latitudinal gradients cause powerful environmental gradients. The topography of alpine areas generates environmental gradients over short distances, and alpine areas are expected to experience greater temperature increase compared to the global average. In this study, we investigate alpha, beta, and gamma diversity, as well as community structure, of vascular plant communities along altitudinal gradients at three latitudes in the Swedish mountains. Species richness and evenness decreased with altitude, but the patterns within the altitudinal gradient varied between sites, including a sudden decrease at high altitude, a monotonic decrease, and a unimodal pattern. However, we did not observe a decline in beta diversity with altitude at all sites, and plant communities at all sites were spatially nested according to some other factors than altitude, such as the availability of water or microtopographic position. Moreover, the observed diversity patterns did not follow the latitudinal gradient. We observed a spatial modularity according to altitude, which was consistent across sites. Our results suggest strong influences of site‐specific factors on plant community composition and that such factors partly may override effects from altitudinal and latitudinal environmental variation. Spatial variation of the observed vascular plant communities appears to have been caused by a combination of processes at multiple spatial scales.     

Morphological variation and floral abnormalities in a trigger plant across a narrow altitudinal gradient

Local adaptation in alpine plants has been demonstrated across wide altitudinal gradients, but has rarely been examined across the alpine‐to‐montane transition that often encompasses only a few hundred metres. Here we characterize morphological variation in leaf and floral characteristics of the trigger plant Stylidium armeria along a narrow altitudinal gradient in the Bogong High Plains in Victoria. Across this gradient, which encompasses the high‐elevation limit of this species, linear changes were found for floral scape height, leaf length and flower number. All these traits decreased with increasing altitude, whereas the frequency of abnormal flowers increased. When plants were grown in a common garden environment, an altitudinal pattern for flower abnormalities was no longer detected. However, altitudinal patterns for leaf length and scape height were maintained, albeit weaker than in the field. This indicates heritable variation for these morphological traits; the altitudinal patterns are likely to reflect the effects of selection by environmental factors that vary with altitude. Selection pressures remain to be identified but have generated both cogradient and countergradient patterns of variation.     

Altitudinal patterns of maximum plant height on the Tibetan Plateau

Aims Several studies have shown that plant height changes along environmental gradients. However, altitudinal patterns of plant height across species are still unclear, especially in regions sensitive to climate change. As canopy height decreases dramatically near the tree line in alpine areas, we hypothesize that plant height across all species also decreases with increasing altitude, and distinct thresholds exist along this gradient.Methods Using a large dataset of maximum plant height and elevation range (400 to 6000 m a.s.l.) of 4295 angiosperms from the regional flora of the Tibetan Plateau, we regressed plant height for every 100 m belt against elevation to explore the relationships. To identify the approximate boundaries where dramatic changes in plant height occurs for herbaceous plants, shrubs, trees, woody plants and all angiosperms, we used piecewise linear regression. Phylogenetically independent contrast was used to test the potential evolutionary influences on altitudinal patterns at the family level.Important findings Results showed that for herbaceous plants, shrubs, trees, woody plants and all angiosperms, plant height decreases significantly as altitude increases. In addition, we found that altitude, a proxy for many environmental factors, had obvious thresholds (breakpoints) dictating patterns of plant height. The results of phylogenetically independent contrast also emphasized the importance of evolutionary history in determining the altitudinal patterns of plant height for some growth forms. Our results highlight the relative intense filtering effect of environmental factors in shaping patterns of functional traits and how this could vary for different ranges of environmental variables.     

高寒草地植物生存策略地理分布特征及其影响因素

植物性状能够反映植物的生存策略,是植物生态学的研究热点之一。植物CSR策略模型将植物物种分为3类:在资源丰富环境中能够最大程度提高生物量的物种(竞争型物种:C策略);在干扰频率较高的环境中能够快速摄取资源并繁殖的物种(投机取巧型物种:R策略);在资源贫瘠环境中能够保持个体生存的物种(耐受型物种:S策略)。植物叶片性状对环境梯度具有适应性的改变,性状的改变对植物生存策略产生影响,但是青藏高原植物叶片性状是如何影响植物CSR生存策略的,其机制尚不清楚。该研究探究了高寒草地植物CSR生存策略的分布特征,以及环境因子对CSR生存策略的影响机制。2020年7–8月,对青藏高原高寒草地53个样点进行了调查,测定植物叶片叶面积、叶片鲜质量和叶片干质量等性状,并计算C、S、R值。然后,分析关键地理环境要素对植物CSR策略影响的主要因子和作用机理。结果表明:(1)在青藏高原高寒草地,植物的生存策略主要以S策略(41.6%–96.7%)为主。(2)随着经度的增加,青藏高原高寒草地C策略植物所占比例自西向东逐渐上升;在海拔梯度上,高寒草地C策略植物所占比例随着海拔的升高而降低。(3)随机森林分析结果显示降水...     

Carbon isotope discrimination by plants follows latitudinal and altitudinal trends

Summary In an earlier paper we provided evidence that carbon isotope discrimination during photosynthesis of terrestrial C3 plants decreases with altitude, and it was found that this was associated with greater carboxylation efficiency at high altitudes. Changing partial pressures of CO₂ and O₂ and changing temperature are possible explanations, since influences of moisture and light were reduced to a minimum by selective sampling. Here we analyse plants sampled using the same criteria, but from high and low altitudes along latitudinal gradients from the equator to the polar ends of plant distribution. These data should permit separation of the pressure and temperature components (Fig. 1). Only leaves of fully sunlit, non-water-stressed, herbaceous C3 plants are compared. The survey covers pressure differences of 400 mbar (ca. 5000 m) and 78 degrees of latitude (ca 25 K of mean temperature of growth period). When habitats of similar low temperature (i.e. high altitude at low latitude and low altitude at polar latitude) are compared, discrimination increases towards the pole (with decreasing altitude and thus increasing atmospheric pressure). Latitudinally decreasing temperature at almost constant atmospheric pressure (samples from low altitude) is associated with a decrease in discrimination. So, polar low-altitude plants have ¹³C values half way between humid tropical lowland and tropical alpine plants. It is unlikely that latitudinal changes of the light regime had an effect, since low and high altitude plants show contrasting latitudinal trends in ¹³C although local altitudinal differences in overall light consumption were small. These results suggest that both temperature and atmospheric pressure are responsible for the altitudinal trends in ¹³C discrimination. Temperature effects may partly be related to increased leaf thickness (within the same leaf type) in cold environments. Theoretical considerations and laboratory experiments suggest that it is the oxygen partial pressure that is responsible for the pressure related change in discrimination. The study also provided results of practical significance for the use of carbon isotope data. Within a community of C3 plants, discrimination in species of similar life form, exposed to similar light, water and ambient CO₂ conditions ranges over 4, with standard deviations for 10–30 species of ±0.6 to 1.2. This natural variation has to be taken into account by using a sufficient sample size and standardization of sampling in any attempt at ecological site characterization using carbon isotope data. Evidence of a pronounced genotypic component of this variation in ¹³C discrimination in wild C3 plant species is provided. Correlations with dry matter partitioning, mesophyll thickness and nitrogen content are also present.     

Seedling establishment of Asteraceae forbs along altitudinal gradients: a comparison of transplant experiments in the native and introduced ranges

Aim Since ecological and evolutionary context changes when a plant species is introduced to a new area, it can be assumed that responses of alien plants to changing conditions along environmental gradients differ from those in their native range. Even if seed availability is not limited, the distribution of alien plants along such a gradient might still be restricted by their ability to germinate and establish as seedlings. In the present study, we aim at testing what factors promote or limit plant invasions during early establishment by using altitude as a model gradient. Location Altitudinal gradients in the Wallowa Mountains (Oregon, USA) and the Swiss Alps (Valais, Switzerland). Methods In transplant experiments along altitudinal gradients, we investigated the early establishment success of eight invasive alien Asteraceae species in their native and introduced ranges in the Wallowa Mountains and the Swiss Alps. Results Seedling recruitment was not restricted to relatively lower altitudes in the introduced range. In addition, we found no evidence for genetic adaptation along the altitudinal gradient in the introduced range, highlighting the importance of phenotypic flexibility for invasions. Furthermore, seedling recruitment was only enhanced by disturbance in the native range where vegetation was comparably dense but not in the introduced range. However, plant development was strongly delayed in the introduced range, probably due to low seasonal water availability. Main Conclusions We conclude that introduced plants, due to their ability to tolerate a wide range of environmental conditions, are not necessarily more restricted in their altitudinal limits than they are in their native range. Furthermore, due to other interacting factors (e.g. different competition situations among ranges), attempts to predict distributions of alien plants in the introduced range that are based on their distributions in the native range may be misleading.     

Experimental measurements of leaf carbon isotope discrimination and gas exchange in the progenies of Plantago depressa and Setaria viridis collected from a wide altitudinal range

Significant correlations between leaf carbon isotope discrimination (Δ) and altitude and between gas exchange and altitude have been reported in previous studies, raising the question of whether the altitudinal variations in discrimination and gas exchange can be attributed to genetic differences among populations from different altitudes. Studies that focus on in situ analysis cannot distinguish the effects of genetic variation from environmental variation. This article describes an experiment in which seeds of Plantago depressa (C3 species) and Setaria viridis (C4 species) collected from a wide altitudinal range were grown in the same environment. Carbon isotopic ratios (δ¹³C) and gas exchange of the seedlings were measured. The progenies of P.   depressa and S.   viridis no longer display any significant Δ decreases with the altitude of origin as seen in situ. Furthermore, photosynthetic rate, stomatal conductance, the ratio of intercellular to ambient CO₂ and intrinsic water use efficiency for P.   depressa and S.   viridis grown in the greenhouse are also not significantly related to the altitude of origin. The observations suggest that altitudinal variations in Δ and gas exchange are not because of genotypic differences, independent of photosynthetic type.     

Divergent patterns of foliar δ13C and δ15N in Quercus aquifolioides with an altitudinal transect on the Tibetan Plateau: an integrated study based on multiple key leaf functional traits

Aims With a close association with plant water availability, foliar δ 13 C had been investigated extensively in alpine regions; however, foliar δ 15 N has rarely been concurrently used as an indicator of plant nitrogen availability. Due to the positive correlations between leaf nitrogen content and foliar δ 13 C and δ 15 N found in previous studies, we expected that they should show consistent patterns along an altitudinal gradient.Methods To test our hypothesis, we measured foliar δ 13 C and δ 15 N in conjunction with multiple key leaf functional traits of Quercus aquifolioides, a dominant species of alpine forest on the eastern slopes of the Sygera Mountains, southeastern Tibetan Plateau from 2500 to 3800 m.Important findings (i) Contrary to our hypothesis, foliar δ 13 C exhibited a significant positive linear relationship with altitude; in contrast, foliar δ 15 N initially increased and subsequently decreased with altitude, the change in trend occurring around 3300 m. (ii) Our analyses indicated that leaf internal resistance and stomatal conductance, rather than photosynthetic capacity indicated by leaf N concentration, apparently explained the altitudinal variation in foliar δ 13 C, while differences in foliar δ 15 N were likely the result of soil N availability. (iii) Principal component analysis revealed a clear association between δ 13 C and a tradeoff between water loss and carbon gain, indicated by traits related to gas exchange such as leaf thickness, density, stomatal properties. In contrast, the second axis was associated with δ 15 N and nitrogen acquisition strategy in Q. aquifolioides across its altitudinal distribution, represented by traits related to nitrogen concentration and stomata per gram of leaf nitrogen.     

Geomorphological disturbance affects ecological driving forces and plant turnover along an altitudinal stress gradient on alpine slopes

The altitudinal gradient is considered as a stress gradient for plant species because the development and fitness of plant communities tend to decrease as a result of the extreme environmental conditions present at high elevations. Abiotic factors are predicted to be the primary filter for species assemblage in high alpine areas, influencing biotic interactions through both competition for resources and positive interactions among species. We hypothesised that the relative importance of the ecological driving forces that affect the biotic interactions within plant communities changes along an elevation gradient on alpine debris slopes. We used multiple gradient analyses of 180 vegetation plots along an altitudinal range from ~1,600 to 2,600 m and single 100 m-bands in the Adamello-Presanella Group (Central Alps) to investigate our hypothesis; we measured multiple environmental variables related to different ecological driving forces. Our results illustrate that resource limitations at higher elevations affect not only the shift from competition to facilitation among species. A geomorphological disturbance regime along alpine slopes favours the resilience of the high-altitude species within topographic/geomorphological traps. An understanding of the ecological driving forces and positive interactions as a function of altitude may clarify the mechanisms underlying plant responses to present and future environmental changes.     

Local adaptation and cogradient selection in the alpine plant, Poa hiemata, along a narrow altitudinal gradient

Alpine environments are particularly susceptible to environmental changes associated with global warming but there is potential for alpine plants to adapt to warming if local adaptation occurs and gene flow allows genotypes adapted to low altitudes to colonize higher altitude sites. Here we examine the adaptive potential of a common alpine grass, Poa hiemata, within the restricted alpine habitat of Australian mountains, across a narrow altitudinal gradient replicated in three areas. Grasses at high altitude sites had shorter leaf lengths and larger circumferences than those at lower sites. Transplant experiments with clonal material and plants grown from seed indicated that these differences were partly genetic, with environmental and genetic factors both contributing to the differences between altitudes. Differences in altitudinal forms were also evident in a common garden experiment. Plants showed a home-site advantage in terms of survival. A fitness analysis indicated that at high altitude sites, selection favored plants with short leaves and larger circumferences, whereas these traits were selected in the opposite direction at the low altitude sites. These findings indicate cogradient selection and potential for both plastic and genotypic shifts in response to climate change in P. hiemata.