卧龙自然保护区油竹子（Fargesia angustissima）生理生态特征的海拔变化

全球气候变化（如温度升高）迅速而强劲地影响着高海拔地区的生态环境,致使植物生长发生变化,因此高山地区成为研究植物环境适应性及其对全球气候变化响应的理想区域。为了认清气候变化如何影响竹子的生长和生理生态特性,本文系统调查研究了卧龙自然保护区内大熊猫典型主食竹种油竹子（Fargesia angustissima（Mitford）T.P.Yi）,从其天然分布下限至上限,在卧龙自然保护区沿海拔梯度研究了分株的比叶面积（SpecificLeaf Area,SLA）、基于单位叶面积和单位叶质量的叶氮含量（Narea,Nmass）,以及组织非结构性碳水化合物（NSC）含量及其组分。研究结果表明,各调查因子对海拔的响应均是非线性的,表现为随着海拔升高,各调查值先降后升,高峰值出现在分布上限区域（1 810 m）,而最低值出现在中海拔区域（1 620 m）,反映了环境因子随海拔的非线性变化。分析认为,油竹子的生长受不同海拔环境影响较大。     

南滚河自然保护区森林土壤酶活性对海拔升高的响应

【目的】研究土壤酶活性对环境因子沿海拔梯度变化的响应,为理解南滚河国家级自然保护区不同海拔水热条件-植被-土壤理化性质与土壤酶活性之间的相互作用过程和机制提供数据支撑。【方法】选取南滚河国家级自然保护区3种不同海拔梯度典型植被带(沟谷雨林、半常绿季雨林和中山湿性常绿阔叶林),研究不同海拔不同土层森林土壤酶活性的变化特征,采用冗余分析和蒙特卡洛检验,分析环境因子沿海拔梯度变化对土壤酶活性的影响。【结果】年均降水量和Simpson指数随海拔升高呈增加趋势,而植物群落多样性(Margalef丰富度指数、Shannon-Wiener多样性指数、Pielou均匀度指数)、年均气温和土壤温度随海拔升高呈下降趋势;土壤有机碳、全氮、全磷、全钾、水解性氮、有效磷和速效钾含量及含水量随海拔升高呈上升趋势,土壤密度和p H随海拔升高呈降低趋势,土壤C/N随海拔升高呈先增后降趋势;土壤脲酶、蔗糖酶、酸性磷酸酶和过氧化氢酶活性随海拔升高呈增加趋势,土壤多酚氧化酶活性随海拔升高呈下降趋势;随土层深度增加,土壤脲酶、蔗糖酶、酸性磷酸酶和多酚氧化酶活性逐渐降低,过氧化氢酶活性逐渐增加;样地微气候、植被和土壤理化性质沿海拔梯度变化对土壤酶活性具有重要影响,年均气温、年均降水量、土壤温度、植物多样性指数和土壤理化性质与土壤酶活性不同程度地相关,单一环境因子对土壤酶活性影响的重要性排序为全磷含量含水量pH有机碳含量年均气温土壤温度水解性氮含量年均降水量全氮含量全钾含量 Margalef丰富度指数土壤密度速效钾含量有效磷含量Shannon-Wiener多样性指数Pielou均匀度指数C/NSimpson指数。【结论】水热条件(年均气温、年均降水量、土壤温度和含水量)、植物多样性和土壤化学性质(p H、有机碳、水解性氮、全氮、全钾和全磷)存在明显的梯度变化,可调控不同海拔梯度内的土壤酶活性,其中土壤磷、水分、p H、碳、氮和钾是调控土壤酶活性海拔响应的关键因素。     

3种海拔对团竹生物量分配和克隆形态特征的影响

通过对四川省北川县片口自然保护区不同海拔梯度下（2700 m、2800 m与2900 m）团竹分株种群特征进行调查，研究其生物量分配格局和克隆形态特征。结果表明，（1）分株及构件生物量随海拔增加而增加；在不同海拔条件下，地下茎生物量分配有显著差异，其他构件生物量分配无明显差异；秆在构件生物量分配比值中最大，为38％；（2）地上构件形态特征随海拔升高而变化显著；地下构件中根的数量、地下茎基茎、地下茎长度和分蘖强度都随海拔升高呈现出先增加后减少的趋势，且在海拔2800 m时达到最大值，因此宜选择海拔2800 m地带的团竹苗用于人工种植。团竹在不同海拔环境中具有生物量分配和克隆形态的可塑性，有利于其种群对水分、光资源等的有效利用。     

高山林线树木光合作用适应性研究进展

高山林线作为一类特殊的生态过渡带, 各类环境因子随海拔升高具有明显的梯度变化规律。林线过渡带物种组成与植被格局变化、林线上下界及树线推移等长期受到生态学与植物学研究的关注。不同生活型的林线植物, 尤其是木本植物对高海拔林线逆境光合响应方式与策略的差异, 直接导致林线植被组成与分布格局的变化, 最终影响林线上下界以及树线的形成。并且, 作为陆地生态系统对全球气候变化响应最为敏感的地区之一, 林线植物光合作用响应性变化特征已成为预测全球气候变化条件下林线植被未来动态变化的重要参数。文中主要从叶片形态与结构以及叶片光合生理代谢等方面, 综述山地木本植物适应林线逆境的相关研究进展, 以期有助于理解高寒地带林线生境植被分布格局形成以及物种组成变化的驱动因子, 为在全球气候变化条件下开展林线植物光合作用响应性研究提供参考作用。     

祁连山林草复合流域降水规律的研究

在祁连山西水试验区排露沟林草复合流域分坡向和海拔设置26个雨量观测点,观测研究了降水变化规律,结果表明:(1)流域内降水具有明显的季节变化,夏秋季降水充沛,冬春季降水稀少;(2)降水变化是大气环流和小气候及环境因子综合作用的结果,月降水量(R,mm)与月水汽压(P,Pha)、气温(T,℃)、空气湿度(W,%)和水面蒸发(E,mm)之间存在显著的线性相关关系:R=-98.206 3 11.735 3P 1.135 0W 0.086 1E-2.294 6T,相关系数为0.948 6;(3)降水量随海拔升高而递增,2 600～3 300 m百米平均递增率为4.95%,当海拔超过3 300 m及接近山顶时,降水出现随海拔升高而下降的趋势;降水空间分布差异除受海拔影响外,还受坡向的影响,阴坡比阳坡降水多7%左右,处在相同海拔和坡向的植被类型差异对降水量的影响不显著;(4)模型Ⅱ能够较准确模拟流域降水动态变化,可以利用流域附近气象站降水观测值进行流域降水量估算.     

云南松生物量和碳储量动态变化分析

基于云南省第五次、第六次和第七次森林资源连续清查样地数据,以优势树种为云南松的样地为研究对象,采用生物量转换因子法及生物量和碳储量之间的转换模型,分别计算和分析不同海拔梯度、龄组、郁闭度、林分密度以及径级的云南松样地生物量和碳储量及其变化情况。结果表明:1)云南松样地生物量和碳储量随海拔的升高呈先升高后降低的变化趋势,1 800~2 400m为云南松生长的最适宜海拔区间;2)随龄组的增加呈先升高后降低的变化趋势,样地生物量和碳储量最高值出现于中龄林;3)随郁闭度的增大呈先升高后降低的变化趋势,最高值出现于中郁闭度林分;4)随林分密度的增加呈先升高后降低的变化趋势,最高值出现于Ⅱ林分密度级;5)随径级的增大呈先升高后降低的变化趋势,最大值出现于中径组。     

轿子山急尖长苞冷杉叶片和气孔特征随海拔梯度变化研究

海拔梯度变化在很小的地理范围内会引起多种环境因子的变化,研究植物性状随海拔梯度的变异特征,对于理解植物对环境适应策略具有重要意义。在云南轿子山国家级自然保护区12个不同海拔(3 510、3 550、3 610、3 660、3 700、3 760、3 790、3 840、3 900、3 960、4 000、4 060 m)样地内观测急尖长苞冷杉的叶片基本特征和气孔特征。结果表明:随海拔的升高,急尖长苞冷杉叶片大小呈双峰型变化,在海拔3 610 m处及4 000 m处存在2个峰值,而叶片厚度无显著变化,气孔密度及大小与海拔存在极小的负相关;综合叶片及气孔特征,海拔变化对急尖长苞冷杉的叶片性状有明显影响,海拔3 610 m为急尖长苞冷杉的最适生长区域;随海拔升高,环境因子逐渐恶劣,逐渐不适应急尖长苞冷杉生长。     

灵石山米槠林优势种不同叶龄叶绿体色素沿海拔梯度的变化

对灵石山米槠林优势种不同叶龄叶绿体色素的差异以及优势种叶绿体色素沿海拔梯度的变化规律进行分析。结果表明:除少数优势种在某海拔梯度以外,不同叶龄叶绿素a、叶绿素b和叶绿素(a+b)存在显著或极显著差异,一般在中高海拔梯度差异最大,且2年生叶片含量高于1年生叶片;而不同叶龄间叶绿素a/b和类胡萝卜素的差异性表现复杂,表现了植物对环境的适应。不同优势种叶绿素[叶绿素a、叶绿素b和叶绿素(a+b)]在海拔梯度间存在显著差异,并且沿海拔梯度的变化呈现一定的规律性,米槠林优势种中有7种优势种不同叶龄叶绿素[叶绿素a、叶绿素b、叶绿素(a+b)]含量均随海拔的升高先升高,一般在A5(442m)或A6(531m)达到最大值,然后随海拔的升高逐渐降低,而叶绿素a/b随海拔升高变化趋势比较多样化。类胡萝卜素作为功能色素由于其功能比较复杂,对海拔的响应规律也不尽相同。海拔作为重要的环境因素对于植物叶绿体色素产生显著影响,不同的优势种沿海拔的变化规律存在差异,说明其对环境的适应对策不同。     

不同经度天山云杉林分因子随海拔梯度的变化

选择新疆天山山脉从西到东处于不同经度的昭苏、巩留、乌苏、乌鲁木齐和哈密的天山云杉林进行垂直样带调查:将各地点划分为低、中和高3个海拔范围,比较不同范围内主要林分因子的差异;分析主要林分因子随海拔梯度(50 m间隔)的变化.结果表明:各地点3个海拔范围内主要林分因子呈现出不同的变化规律;天山云杉林分平均胸径和最大胸径随海拔梯度的变化差异不显著,而平均树高、最大树高、林分密度、胸高断面积和蓄积量随海拔的增加总体上呈现单峰型的变化趋势;尽管不同经度位置天山云杉林分布的海拔上下限范围不同,但各个位置天山云杉林分的平均树高、最大树高、林分密度、胸高断面积和蓄积量随海拔梯度的变化都可以用二次曲线方程进行描述.     

长白山沿海拔梯度臭冷杉径向生长对气候变化的响应

为了研究树木生长对气候变化的响应,选取长白山北坡3个海拔高度建立了臭冷杉Abies nephrolepis的年轮差值年表。通过年表特征及响应函数分析,探讨不同海拔臭冷杉径向生长对气候变化的响应。结果表明,温度是影响海拔分布上限(1 760 m)臭冷杉生长的主要影响因子,年轮宽度指数与上年9月和当年5月的平均温呈显著正相关,与降水没有显著关系;在中海拔(1 250 m),年轮宽度指数与温度和降水都没有呈现出显著相关关系;在海拔分布下限(780 m),降水是主要的影响因子,年轮宽度指数与上年9月和10月的降水量呈显著正相关,与温度没有达到显著相关。未来气候变化有利于臭冷杉的径向生长,有利于长白山暗针叶林生产力的增加。     

西双版纳兰科植物海拔分布格局

根据实地调查和相关文献资料整理,确定西双版纳野生兰科植物428种。将研究区按200 m海拔段划分为10个海拔梯度,分析兰科植物垂直分布格局。结果表明:①西双版纳兰科植物的垂直分布格局呈"中间膨胀型",峰值出现在1 000~1 200 m海拔段;②相邻海拔段的兰科植物种类相似系数较高(0.31~0.79),海拔段相隔越远相似性系数越小;③随海拔升高各海拔段的兰科植物生活型及区系成分的种类数量也呈现出先升高后下降的垂直分布格局;④地生兰和附生兰的种数在海拔600 m左右达到平衡,600 m以上附生兰占优势;⑤各海拔段兰科植物均以热带种类占主导地位(72%~87.8%),其中又以热带亚洲分布型的种类占优势。     

Comparison of fractal characteristics of species richness patterns among different plant taxonomic groups along an altitudinal gradient

This study was done using the non brown fractal model to quantify and compare the variations in the species richness of trees, shrubs, herbs and all plants along an altitudinal gradient and to characterize the dominating ecological processes that determine the variations. Two transects were sampled far away from any anthropogenic disturbances along the shady slopes of the Dongling mountains in Beijing, China. Both transects were continuous and 2 m wide, and every individual tree and shrub was recorded in each of them. Discrete quadrats of 1 m × 1 m were located along the transects A and B for estimation of the herb species richness along the altitudinal gradients. The level interval between the quadrats was 10 m and 25 m respectively. In this study, transects A and B were combined into one transect AB, and 40 m was selected as the optimal quadrat length along the altitudinal gradients for measuring the plant species richness patterns. Species richness in each quadrat was calculated using a program written in Matlab 6.0. Direct gradient analysis was used to describe the overall trends in the species richness of trees, shrubs, herbs and other plants with change in altitude, while the non-brown fractal model was used to detect more accurately their variations at various scales along the gradient. The model assumed that each class of ecological processes affecting the distribution of a variable could be represented by an independent spatial random function. Generally, ecological phenomena are determined not by a single ecological process but by multiple ones. These processes act on ecological patterns within their own spatial scales. In the non-brown fractal model, the spatial random functions are nested within a larger range of spatial scales. The relative contribution of the spatial random functions to the spatial variation of a variable is indicated by a weighting parameter that has to be greater than or equal to zero. In this paper, we reached the following results and conclusions. Firstly, the direct gradient method describes the general trends of trees, shrubs, herbs and all plants along the altitudinal gradient but is unable to provide further details on the altitudinal variations in the species richness. The non-brown fractal model brought out the altitudinal variations in the species richness of trees, shrubs and herbs at various scales and related them to the ecological processes. The sharp changes in the double-log variograms suggest that the non-brown fractal model is suitable for characterizing the altitudinal patterns in the species richness of trees, shrubs and herbs at various scales but is not appropriate for explaining the variations in the plant species richness, since no significant changes were found in the double-log variograms in this case. Secondly, for the trees, the double-log variogram was divided into two scale ranges (0–245 m and 245–570 m), with a fractal dimension of 1.83 and 1.10, respectively, implying that changes in the tree species richness were random at small scales (0–245 m) and almost linear at large scales (245–570 m) along the altitudinal gradients. This suggests that altitudinal variations in the tree species richness are dominated by short-range processes at small scales and by long-range processes at large scales. Thirdly, for shrubs and herbs, the double-log variograms exhibited three ranges (0–101 m, 125–298 m and 325–570 m), and the fractal dimensions were 1.78 and 1.97, 1.56 and 1.43, and 1.08 and 1.25, respectively. The results indicate that, as in the case of trees, species richness of shrubs and herbs are distributed randomly at small scales and change in a linear manner at large scales although variations in the herb species richness is less heterogeneous than shrub species richness at large scales. These results also indicate that species richness of shrubs and herbs change approximately like brown movement at middle scales. The results also suggest that altitudinal variations in the specie richness of shrubs and herbs are dominated by three ecological processes, short-range ecological processes at small scales, long-range ecological processes at large scales, and brown fractal processes at middle scales. Interestingly, comparisons of the variations in the species richness of shrubs and herbs reveal that shrubs and herbs present the same scale range in spatial variation in species richness but display different trends in species richness along the altitudinal gradient, i.e. the shrub species richness decreased with increasing elevation whereas the herb species richness peaked at the mid-high elevation. These patterns suggest that although the scales at which the main processes affect patterns in species richness are the same, the processes are completely different, or the processes are similar but the responses of the shrubs and herbs to the ecological processes are different. Finally, the plant species richness did not show any obvious pattern along the altitude gradient and maintained a constant fractal dimension across all scales, this is perhaps because the processes defining the patterns of plant species richness had similar weights and acted over closely related scales. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(6): 901–909 [译自: 植物生态学报]     

Larix olgensis growth-climate response between lower and upper elevation limits:an intensive study along the eastern slope of the Changbai Mountains,northeastern China

Larix olgensis is a dominant tree species in the forest ecosystems of the Changbai Mountains of northeast China.To assess the growth response of this species to global climate change,we developed three tree-ring width and biomass chronologies across a range of elevations in the subalpine forests on the eastern slope of the Changbai Mountains.We used dendroclimatic analyses to study key factors limiting radial growth in L.olgensis and its variation with elevation.The statistical characteristics of chronologies suggested that elevation is a determinant of tree growth patterns in the study area.Response function analysis of chronologies with climate factors indicated that climate–growth relationships changed with increasing elevation:tree growth at high elevation was strongly limited by June temperatures of the previous year,and as elevation decreases,the importance of temperature decreased;tree radial growth at mid-elevation was mainly controlled by precipitation towards the end of the growing season of the current year.Biomass chronologies reflected a stronger climatic signal than tree-ring width chronologies.Spatial correlation with gridded climate data revealed that our chronologies contained a strong regional temperature signal for northeast China.Trees growing below timberline appeared to be more sensitive to climate,thus optimal sites for examining growth trends as a function of climate variation are considered to be just below timberline.Our study objective was to provide information for more accurate prediction of the growth response of L.olgensis to future climate change on the eastern slope of the Changbai Mountains,and to provide information for future climate reconstructions using this tree species in humid and semi humid regions.     

岷江上游干旱河谷白刺花生物量及其与土壤含水量关系研究

岷江上游干旱河谷区是我国长江上游的主要生态功能服务区,而该区域由于“受焚风效应”的影响以及频繁的滑坡、泥石流等地质灾害影响,使该区域也是我国生态环境十分脆弱区。本文以该区域内分布最广泛的灌木——白刺花为对象,研究其生物量、生物量分配格局以及生物量分配与土壤含水量的关系,为干旱河谷区灌丛植被的保护和恢复重建提供理论依据。研究结果表明：①白刺花的地上生物量分配中,阴坡叶片所占地上生物量比例变化范围在36．01％-41．54％,明显高于阳坡（14．35％～21．12％）;AK海拔梯度变化上看,白刺花在阳坡与阴坡的茎枝、叶和种子各部分生物量以及单丛生物量大小变化趋势均呈现“V”字型变化趋势。②白刺花单丛地下生物量则阴坡海拔1860m地最高,达到440．06g,阳坡海拔2280m最高,为436．39g;最低的地下生物量主要在阴坡海拔1730m和阳坡海拔1880m,仅为47．23g和44．61g,最高生物量分别是它们的9．9、9．8和9．8、9．2倍。同时,在阴坡和阳坡的不同海拔梯度上,均表现出粗根和中根生物量占地下部分总生物量50％以上,细根、小根生物量之和在30％以下。③白刺花地上各部分生物量与土壤含水量间相关关系显著,并随土壤含水量的增加而增加。其中白刺花单丛地上部分生物量和枝（干）生物量与土壤含水量的回归曲线方程模型Y=27．581x^1.1473和Y=18．648e^0.3454x。最优,其群分别达到0．9677、0．9464;叶生物量和种子生物量与土壤含水量的回归模型则分别以Y=7．3936x^0.3454x和Y=2．5705e^0.4144x最佳,砰分别为0．7172、0．7446。白刺花各部分根系生物量与土壤含水量的变化趋势总体上也相对一致,即随着含水量的增加而增加。但各径级根系生物量与土壤含水量的回归拟合效果相对地上生物量较差,其中粗根生物量?     

北川不同海拔珙桐种群年龄结构研究

采用树木年代学的方法,按照珙桐的分布特点分海拔下限（1400 m～1500 m）,中部（1 600 m～1 700 m）,上限（1 800 m～1 900 m）,研究不同海拔梯度下北川珙桐种群的年龄结构,结果表明：海拔上限珙桐年龄结构偏大,缺乏幼苗及30 a～40a的个体;在海拔中部,30 a～40 a的个体占的比例较大,争夺空间和资源的能力更强;海拔下限的实生幼苗较多.在中部海拔分布的珙桐较多,且年龄结构集中在30 a～40 a之间,低海拔的种群特征是以幼苗为优势,而高海拔面有的幼苗却很少,表明海拔对珙桐种群年龄结构有重要的影响.     

Vertical patterns of the flora of seed plants in Dawei Mountain in Yunnan Province, Southwest China

Vertical gradients incorporate multiple resources gradients which vary continuously. Therefore, research on mountain flo-ristic patterns along vertical gradients is important to reveal regular patterns of the flora along environmental gradients and to under-stand the changes in biodiversity along these gradients and their biological fitness. This study was designed to explore the character-istics of the floral compositions and ecological significance of floristic patterns along the vertical gradients of the National Nature Reserve of Dawei Mountain, located in the southeast of Yunnan Province. We analyzed the structural characteristics of the flora and the distribution patterns of its floristic components as a function of elevation on the basis of our field investigations along vertical vegetation transects. We carried out a systematic cluster analysis in order to determine the dividing line of floristic changes by eleva-tion along gradients and studied the effects of mountain climate on the vertical variation of floristic composition. The study shows: 1) that the obvious boundary, which differentiates tropical distribution, is located at an elevation of approximately 1,500 m, which separates the tropical rain forests from the evergreen broad-leaved forests; 2) that humid rain forests are found below 700 m elevation, mountain rain forests between 700 and 1,500 m, monsoon evergreen broad-leaved forests between 1,300 and 1,800 m and mountain mossy evergreen broad-leaved forests above 1,800 m. Non-representative mountain mossy dwarf forests (above 2,100 m) in the area are found on the windward sides and barren lands on mountain slopes; 3) that Hopea mollissima is one of the major component species of mountain rain forests, but it should not be considered as the major indicator species in humid rain forests as is generally accepted.     

Altitudinal range shift detected through seedling survival of Ceiba aesculifolia in an area under the influence of an urban heat island

Changes in the position along the elevational gradient for plant species are a predicted consequence of global and local climate change. Within the area of influence of cities it is necessary, for conservation and ecological restoration, to understand the effect on plant communities of climate change and urban heat islands, that can increase the temperature around cities as much as 8 °C when compared with peri-urban rural areas. We explored patterns of seedling survival of Ceiba aesculifolia, a tropical tree species, along an altitudinal gradient in a slope facing the city of Morelia, in Michoacán, México, because the city has followed a trend of increasing mean annual temperatures with a difference of at least 4 °C when compared with the surrounding rural areas. The highest survival was found between 2200 and 2230 m a.s.l. (100% after 1 year of planting and 75% after 2 years), 160 m higher than the altitudinal limit of the remnant adult trees in the study area, and close to the highest limit reported for the species at a regional scale (2200 m a.s.l.). There was a significant effect of altitude on survival among experimental plots (P < 0.0001) and the interaction between elevation and distance from the north side of the experimental area was significant (P = 0.017). For restoring populations of C. aesculifolia within our study region, assisted migration through the establishment of populations at the elevational limit or higher than its historical range might be necessary in areas close to cities that already are under the effects of increased temperatures, but might be necessary also in rural areas for ameliorating the expected effects of global climate change in tropical rural areas.     

Drought sensitivity of Norway spruce is higher than that of silver fir along an altitudinal gradient in southwestern Germany

Context

For Central Europe, climate projections foresee an increase in temperature combined with decreasing summer precipitation, resulting in drier conditions during the growing season. This might negatively affect forest growth, especially at sites that are already water-limited, i.e., at low elevation. At higher altitudes trees might profit from increasing temperatures.

Aims

We analyzed variations in radial growth of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) along an altitudinal gradient from 400 until 1,140 m a.s.l. in the Black Forest, to assess climate responses with increasing elevation.

Methods

Climate–growth relationships were analyzed retrospectively using tree-ring and climate data. In total, we sampled stem discs of 135 trees to build 27 species- and site-specific chronologies (n _fir?=?13, n _spruce?=?14).

Results

Our results indicate distinct differences in climate–growth relations between fir and spruce along the gradient. Growth of high-altitude fir was positively related to temperature from January till March. Growth of low-altitude fir and spruce at all elevations was positively related to precipitation and negatively to temperature during the growing season, particularly in July. A self-calibrating Palmer drought severity index underlined summer drought sensitivity of these trees.

Conclusion

Overall, we found that climatic control of tree growth changes over altitude for fir. For spruce, a remarkable synchrony in growth variation and climate response was shown, which indicates that this species is drought sensitive at all studied elevations. In a future warmer climate, the growth of low-altitude fir and spruce along the entire studied gradient may be negatively affected in the Black Forest, if an increased evaporative demand cannot be compensated by increased water supply.