川西亚高山桦木林的林地水文效应

川西亚高山森林是我国西南亚高山林区水源涵养林的重要组成部分,原生的亚高山暗针叶林在经历大规模采伐利用后,天然更新的次生桦木林已成为该区域的主要森林类型之一。前人对原始暗针叶林水文学的研究已相当丰富,内容涉及冠层截留、地被物持水特征、森林蒸发散、土壤入渗、根土作用层等诸多方面;而对于采伐后人工林和天然次生林的研究较少,仅有的结论也以人工林为主。通过对林地苔藓、枯落物和土壤的野外调查与室内实验,分析了川西亚高山次生桦木林在不同林龄和海拔梯度间的林地水文效应,这对于丰富亚高山森林水文学的研究、确定长江上游水源涵养林的恢复与重建模式,都具有重要的意义。研究表明:桦木林苔藓蓄积量及最大持水量在不同林龄间差异显著,随林龄增大而显著增加;而在不同海拔间差异不显著。枯落物蓄积量及最大持水量在不同林龄及海拔间均差异显著,随林龄的增大而增加;在林龄相同的条件下,在中海拔(3200m、3400m)较高,在较高(3600m)、较低(3000m)海拔偏低。苔藓最大持水率平均为945%,在林龄和海拔间差异不显著;枯落物最大持水率平均573%,在林龄和海拔间均差异显著。各林龄和海拔梯度上的桦木林,随土壤深度的增加土壤容重均显著增大,最大持水量显著下降,但毛管持水量和最小持水量仅在部分类型下降显著。土壤0~40cm最大持水量在不同林龄间差异不显著,而在不同海拔间差异显著;这种差异主要表现在林龄10~25a的林分,随海拔升高土壤0~40cm最大持水量增大。在大规模采伐后,苔藓层的恢复是一个长期过程,可以作为次生林地水文效应向原始暗针叶林恢复程度的一个指标。     

马占相思（Acacia mangium）也湿地松（Pinus elliotii）人工林枯落物层的水文生态功能

分析了马占相思与湿地松人工林枯落物的蓄积量、年凋落量及凋落动态、枯落物层对大气降水的截留、以及枯落物抑制土壤水分蒸发和阻滞径流的效应。结果表明：①15龄的马占相思林枯落物蓄积量32.3t/hm^2,年凋落量11.14t/hm^2,最大持水率253.7％,最大持水量28.26t/hm^2;15龄的湿地松林枯落物蓄积量18.7t/hm^2,年凋落量7.30t/hm^2,最大持水率216.7%,量大持水量15.82hm^2;②2种林分对大气降水的截留率分别为15.9%和11.7%,截留率随1次降水降水量（＞10mm）的增加而减少;③2-4cm枯落物覆盖下不同含水量的土壤水分蒸发比无覆盖的土壤减少18.2%-78.3%,枯落物层减少土壤水分蒸发的效应随枯落物层厚度和土壤含水量的增大而增加;④2种枯落物对径流流出时间的阻滞效应随径流深（＜3mm）和坡度的增加而减小,随枯落物层厚度的增加呈直线增加。通过与部分其它森林类型枯落物层水文生态功能比较,认为马占相思与湿地松林枯落物层具有较为优越的水文生态功能。     

川西亚高山林区不同土地利用与土地覆盖的地被物及土壤持水特征

川西亚高山森林是我国西南亚高山水源涵养林的重要组成部分.随着20世纪中叶以来农业人口的增加和森林的开发利用,受干扰强度、频度和干扰时间的影响,原始暗针叶林退化为耕地、草地、灌丛、次生阔叶林或人工林.1998年后,该区相继启动天然林保护工程和退耕还林工程.为评价工程效益,确定长江上游水源涵养林的恢复与重建模式,需要进行不同土地利用类型之间生态水文效应的对比分析.通过野外调查与室内实验,对比分析了川西亚高山林区农田、草地、退耕还林地、灌丛、次生桦木林、人工云杉林和老龄针叶林的地被物(苔藓与枯落物)和土壤持水特征,结果表明:不同土地利用与覆盖类型间地被物和土壤持水性能差异显著.随着干扰程度的增加,苔藓、枯落物蓄积量及最大持水量下降,土壤容重增加,土壤持水性能下降.苔藓最大持水量排序是老龄针叶林>人工云杉林>天然次生林>灌丛.枯落物最大持水量排序是老龄针叶林>天然次生林>人工云杉林>灌丛>草地>退耕还林地.人工云杉林与天然次生林之间、草地与退耕还林地之间苔藓和枯落物最大持水量没有显著差异.土壤0～40cm最大持水量排序是天然次生林>老龄针叶林>人工云杉林>灌丛>农田>草地>退耕还林地,其中天然次生林显著高于人工云杉林,草地与退耕还林地之间没有显著差异.对于森林恢复途径和树种的选择,需要考虑未来林分的多种生态系统服务功能.     

西双版纳热带山地雨林枯落物及其土壤水文功能

研究了云南西双版纳热带不同海拔梯度山地雨林枯落物层及土壤层水文功能.结果表明: 土壤容重随着海拔的增加而降低,土壤总孔隙度、非毛管孔隙度、毛管孔隙度、土壤最大持水率、最大持水量、有效持水量和土壤含水量随海拔的增加而增加,局部有所波动;雨季前期含水量、饱和含水量和有效调蓄水空间随海拔的增加而增加,其中,饱和含水量和土壤有效调蓄水空间在不同海拔区差异均显著(P<0.05).土壤渗透性能与总孔隙度和非毛管孔隙度均呈极显著正相关关系(P<0.01),其中,非毛管孔隙对土壤渗透性的影响更为显著.不同海拔枯落物未分解层厚度均占总厚度的一半以上,枯落物厚度均表现为未分解层＞半分解层;枯落物总蓄积量和半分解层蓄积量占枯落物总蓄积量的比例均随海拔的增加而增加,说明低海拔枯落物分解速度较慢,高海拔枯落物分解速度较快.不同海拔枯落物半分解层和未分解层最大持水量、最大持水率、自然含水率、有效拦蓄率和有效拦蓄量均随海拔的增加而增加,并且各海拔未分解层均高于半分解层,而有效拦蓄量深度随海拔的增加而降低,局部有所波动.综合未分解层和半分解层的变化规律可知,高海拔拦蓄能力较强,低海拔较弱.不同海拔枯落物持水量随着浸泡时间增加而增加;枯落物吸水速率随着浸泡时间增加而降低,12 h后枯落物吸水速率逐渐趋于饱和.不同海拔枯落物持水量与浸水时间可用对数方程表示;吸水速率与浸泡时间可用冥函数方程表示.综合分析各项因子,低海拔热带山地雨林水源涵养能力普遍低于高海拔.     

马占相思 (Acacia mangium)与湿地松 (Pinus elliotii)人工林枯落物层的水文生态功能

分析了马占相思与湿地松人工林枯落物的蓄积量、年凋落量及凋落动态、枯落物层对大气降水的截留、以及枯落物抑制土壤水分蒸发和阻滞径流的效应.结果表明①15龄的马占相思林枯落物蓄积量32.3t/hm2,年凋落量11.14t/hm2,最大持水率253.7%,最大持水量28.26t/hm2;15龄的湿地松林枯落物蓄积量18.7t/hm2,年凋落量7.30t/hm2,最大持水率216.7%,最大持水量15.82hm2;②2种林分对大气降水的截留率分别为15.9%和11.7%,截留率随1次降水降水量(>10mm)的增加而减少;③2～4cm枯落物覆盖下不同含水量的土壤水分蒸发比无覆盖的土壤减少18.2%～78.3%,枯落物层减少土壤水分蒸发的效应随枯落物层厚度和土壤含水量的增大而增加;④2种枯落物对径流流出时间的阻滞效应随径流深(<3mm)和坡度的增加而减小,随枯落物层厚度的增加呈直线增加.通过与部分其它森林类型枯落物层水文生态功能比较,认为马占相思与湿地松林枯落物层具有较为优越的水文生态功能.     

不同经营模式对川西亚高山天然次生林林地水文效应的影响

天然次生林是川西亚高山林区经历大规模砍伐后形成的主要森林类型之一,是我国西南林区水源涵养林的重要组成部分。以不同经营模式(抚育经营、清林+补植经营以及封育经营)的川西亚高山次生桦木林和桦木、岷江冷杉混交林为研究对象,通过样方取样法获取和分析了林地苔藓、枯落物和土壤的水文指标。结果表明,与封山育林经营相比,抚育经营下的两种林型的苔藓最大持水率均显著升高(F=8.147,P=0.010;F=15.525,P=0.006)、桦木林的蓄积量显著降低(F=4.979,P=0.022),而苔藓最大持水量变化不显著;混交林则均无显著变化。在清林+补植经营下,混交林苔藓水文效应变化不显著(F=2.280,P=0.183),而桦木林虽然苔藓最大持水率无显著变化(F=4.072,P=0.098),但蓄积量的显著降低(F=3.536,P=0.044)导致了其最大持水量的降低(F=3.782,P=0.042)。两种经营方式基本上促进了天然林的枯落物最大持水率、降低了林下枯落物蓄积量;其中抚育经营效果更显著,但两种经营方式下枯落物最大持水量变化不显著。两种经营方式下,桦木林和混交林的林下土壤容重均降低(F=10.715,P0.01;F=5.148,P0.05),同时桦木林土壤最大持水量增加(F=4.499,P0.05),其中抚育经营的影响程度都更显著。从4年来的短期效应来看,两种经营方式均对天然林的林地持水能力具有促进作用,抚育经营较清林+补植经营更显著,但这仅是短期的结果,两种经营方式对于退化天然林水文以及其他生态功能恢复的长期影响还有待于进一步的观测研究。     

松嫩草原三种主要植物群落枯落物层生态水文功能

分析了松嫩草原主要植物群落羊草群落、虎尾草群落、碱茅群落枯落物层的蓄积量及持水能力、枯落物层对降水的截留以及枯落物层抑制土壤水分蒸发的效应.结果表明,羊草群落枯落物蓄积量最大为4.7t·hm-2,最大持水量为9.6t·hm-2,最大持水率为208.4%;虎尾草群落枯落物层蓄积量、最大持水量和最大持水率分别为3.0t·hm-2、7.4t·hm-2和262.8%;碱茅群落分别为2.6t·hm-2、5.0t·hm-2和202.2%;3种群落枯落物层对降水的截留量分别为6.57、5.79和5.26t·hm-2,随着降雨量的增加,截留量增加,截留率减小;0.5~2mm枯落物覆盖下不同含水量的土壤水分蒸发比无覆盖的土壤减少7.95%~56.79%,枯落物层减少土壤水分蒸发的效应随枯落物层厚度和土壤含水量的增大而增加.     

长白山暗针叶林苔藓枯落物层的降雨截留过程

对长白山北坡暗针叶林地面苔藓枯落物层 (苔藓植物和枯落叶交织而成的层状结构 )持水能力、截留降雨过程及动态变化进行了研究。结果表明苔藓枯落物层持水能力很强 ,最大持水量相当于 4 .8mm的降雨量 ,这个结果大于将苔藓与枯落物分开来测定的结果之和 ,说明成层结构能够提高持水能力。在一次降雨过程中对降雨截留过程的观测发现 :持水量刚开始随降雨增加而增加 ,到一定量时 ,降雨再增加 ,持水量反而缓慢下降。对持水动态变化一个月的观测 ,表明苔藓枯落物层水分蒸发较快 ,当长时间无雨时 ,持水量显著下降 ,当降雨再次发生时 ,持水量迅速回升 ,这种动态过程伴随着降雨的发生而不断往复。此外 ,在夜间苔藓枯落物层有吸收空气中水气的特点 ,吸收量受空气湿度的影响     

杨麦间作系统枯落物持水能力对地表径流氮流失的影响

以太湖流域杨麦间作系统为研究对象, 对系统内不同枯落物蓄积量持水能力及对地表径流中不同形态氮流失效应进行定位观测, 结果表明: 随着枯落物蓄积量的增加, 蓄积量为0.4 t·hm–2最大持水量、最大拦截量和有效拦截能力＞0.2 t·hm–2＞0.1 t·hm–2, 最大持水量的变化范围为2.35-12.93 t·hm–2, 最大拦截量变化范围2.17-10.95 t·hm–2, 有效拦蓄能力在1.80-9.10 t·hm–2之间; 枯落物持水量与浸泡时间呈对数关系(R2＞0.90), 枯落物吸水速率与浸泡时间呈幂函数关系(R2＞0.99)。枯落物蓄积量越大, 对地表径流量及地表径流中不同形态氮流失的削减效果越显著。而2种间作密度下, A1处理对地表径流中TN、NO3–-N和NH4+-N的削减效果＞A2; 径流中以NO3–-N流失为主, TN、NO3–-N和NH4+-N流失量随枯落物蓄积量的增加呈下降的趋势, 蓄积量越大, 差异越显著。      

辽东低山区5种典型水源涵养林枯落物持水特性

对辽东低山区5种水源涵养林枯落物持水特性进行研究,可以为该区水源涵养林的营造、科学经营提供理论依据。2018年6月在辽宁省抚顺县国有温道林场选取纯林(落叶松林(Larix olgensis)、油松林(Pinus tabuliformis)、红松林(Pinus koraiensis)、刺槐林(Robinia pseudoacacia)和杂木林为研究对象,调查各林分枯落物厚度、蓄积量等,并用浸泡法测定最大持水量、最大持水率,建立持水量、吸水速率与浸水时间之间的关系。结果表明:(1)5种林分枯落物厚度3.6～7.8 cm,平均厚度6.2 cm;蓄积量15.40～50.38 t·hm-2,平均值30.42 t·hm-2。(2)枯落物最大持水量13.61～27.21 t·hm-2,大小排序为杂木林落叶松林刺槐林红松林油松林;最大持水率变化稍有不同,依次为刺槐林落叶松林杂木林红松林油松林。(3)枯落物有效拦蓄量19.60～142.67 t·hm-2,大小排序为落叶松林红松林杂木林刺槐林油松林。(4)回归分析表明,枯落物持水量与浸水时间符合关系式Q=alnt+b,相关系数R2均大于0.80;吸水速率与浸水时间符合关系式V=ctn,相关系数R2均大于0.99。综上,落叶松林、红松林枯落物蓄积量最大、持水能力和有效拦蓄能力均较强,刺槐林、杂木林次之,油松林较差。     

UV-B-tolerance of alpine and arctic Daphnia

Survival of transparent and pigmented Daphnia from alpine as well as arctic habitats, exposed to solar radiation with total or reduced UV-B, was measured. In the alpine pond, survivorship and reproduction of transparent juvenile as well as adult Daphnia were significantly reduced due to UV-B. Transparent adult arctic Daphnia also showed significant reduction in survivorship when exposed to total, when compared with controlled solar radiation. In contrast, heavily melanized adult arctic Daphnia showed no increase in mortality when exposed to natural solar radiation. There appeared to be differences both in tolerance and behavior in the two arctic forms when exposed to solar radiation.     

Ecology of alpine streams

1. This review examines ecological conditions and zoobenthic communities of kryal, krenal and rhithral streams of the alpine zone. Altitudinal and biogeographical faunal patterns are also analysed. 2. Kryal segments, fed by glacial meltwater, are characterized by low temperatures (T_max4°C) and large diel flow fluctuations in summer. The water may be clear or turbid from suspended rock flour. Fishes and higher plants are absent. The macroalga Hydrurus foetidus may be abundant in kryal and other alpine stream types of the Holarctic. The highly restricted cosmopolitan fauna of glacial brooks consists of diamesine chironomids, sometimes accompanied by simuliids. Sparse food resources include algae and allochthonous (aeolian) organic matter. 3. Rhithral segments in alpine catchments are characterized by soft water, a hydrograph dominated by an extended period of snowmelt runoff, and a broader temperature range than kryal or krenal biotopes. Bryophytes, macroalgae (chrysophytes, chlorophytes, cyanophytes, rhodophytes) and epiphytic and epilithic diatoms constitute the flora. A relatively diverse zoobenthos includes four orders of insects (Flecoptera, Ephemeroptera, Trichoptera, Diptera), turbellarians, acarines, oligochaetes and nematodes. 4. Krenal segments, fed by groundwater, are typically calcareous with summer-cool and winter-warm thermal conditions, high water clarity, and constant flow regimes. Bryophytes and macroafgae are accompanied by a rich diatom flora. The zoobenthos consists of a composite of kryal and rhithral elements with few crenobionts. Zoobenthos species richness values are intermediate between those of kryal and rhithral segments, whereas densities in perennial, well-oxygenated springbrooks far exceed those in other alpine stream types. 5. Downstream faunal changes are most predictable in kryal segments where chironomids of the genus Diamesa are the predominant, if not sole, members of the zoobenthos in the upper zone of glacial brooks, the metakryal. Where T_max exceeds about 2°C the transition to the hypokryal occurs and Diamesa is co-dominant with simuliids. These largely stenozonal headwater forms decline downstream where T_max exceeds about 4°C, concomitant with a marked increase in the euryzonal mountain fauna. 6. Species occurring in alpine rhithral biotopes tend to be euryzonal forms at their upper alrirudinal limits, whereas the lower elevation mountain stream fauna consists of species with narrower distribution limits. There is, however, a precipitous drop in mean altitudinal range from the alpine rhithral to the kryat because of the stenozonal nature of the glacial brook fauna. 7. The view that effects of temperature on generation time and mutation rate determine the speed at which selection proceeds is consistent with altitudinal species richness patterns exhibited by zoobenthos along the alrirudinal gradient and may provide an evolutionary explanation for the low faunal diversity in alpine headwaters. 8. With increasing altitude, mountain ‘islands’ become progressively insular as area decreases and isolation increases. For a cold-adapted stream fauna the insular nature of mountain tops is greatest in the tropics. Nonetheless, alpine stream faunas generally     

Dormancy of alpine and subalpine perennial forbs

Depth of dormancy of alpine and subalpine perennial forbs in autumn was investigated, which was judged by the number of days required for growth initiation at 24 °C. The depth of dormancy differed depending on Raunkiaer’s life-form and shoot habits. Chamaephytes with perennial shoot-axes showed shallower dormancy than hemicryptophytes with annual shoot-axes, and geophytes with annual shoot-axes showed the deepest dormancy. The results strongly suggest that the dormancy is more endogenously controlled in forbs less hardy to freezing stress. Potential growth ability of alpine herbaceous chamaephytes in autumn is an adaptive advantage, since they utilize the short vegetative period as long as possible. All of the species with annual shoot-axes had winter buds covered with scales. In plants with perennial shoot-axes, percentage of winter buds covered with scales increased with increasing depth of dormancy. The results indicate that the shoot apices are well protected by bud scales in forbs with a long endogeneous-dormant period.     

Functional traits and root morphology of alpine plants

Background and Aims

Vegetation has long been recognized to protect the soil from erosion. Understanding species differences in root morphology and functional traits is an important step to assess which species and species mixtures may provide erosion control. Furthermore, extending classification of plant functional types towards root traits may be a useful procedure in understanding important root functions.

Methods

In this study, pioneer data on traits of alpine plant species, i.e. plant height and shoot biomass, root depth, horizontal root spreading, root length, diameter, tensile strength, plant age and root biomass, from a disturbed site in the Swiss Alps are presented. The applicability of three classifications of plant functional types (PFTs), i.e. life form, growth form and root type, was examined for above- and below-ground plant traits.

Key Results

Plant traits differed considerably among species even of the same life form, e.g. in the case of total root length by more than two orders of magnitude. Within the same root diameter, species differed significantly in tensile strength: some species (Geum reptans and Luzula spicata) had roots more than twice as strong as those of other species. Species of different life forms provided different root functions (e.g. root depth and horizontal root spreading) that may be important for soil physical processes. All classifications of PFTs were helpful to categorize plant traits; however, the PFTs according to root type explained total root length far better than the other PFTs.

Conclusions

The results of the study illustrate the remarkable differences between root traits of alpine plants, some of which cannot be assessed from simple morphological inspection, e.g. tensile strength. PFT classification based on root traits seems useful to categorize plant traits, even though some patterns are better explained at the individual species level.     

Global distribution and bioclimatic characterization of alpine biomes

Although there is a general consensus on the distribution and ecological features of terrestrial biomes, the allocation of alpine ecosystems in the global biogeographic system is still unclear. Here, we delineate a global map of alpine areas above the treeline by modelling regional treeline elevation at 30 m resolution, using global forest cover data and quantile regression. We then used global datasets to 1) assess the climatic characteristics of alpine ecosystems using principal component analysis, 2) define bioclimatic groups by an optimized cluster analysis and 3) evaluate patterns of primary productivity based on the normalized difference vegetation index. As defined here, alpine biomes cover 3.56 Mkm² or 2.64% of land outside Antarctica. Despite temperature differences across latitude, these ecosystems converge below a sharp threshold of 5.9°C and towards the colder end of the global climatic space. Below that temperature threshold, alpine ecosystems are influenced by a latitudinal gradient of mean annual temperature and they are climatically differentiated by seasonality and continentality. This gradient delineates a climatic envelope of global alpine biomes around temperate, boreal and tundra biomes as defined in Whittaker's scheme. Although alpine biomes are similarly dominated by poorly vegetated areas, world ecoregions show strong differences in the productivity of their alpine belt irrespectively of major climate zones. These results suggest that vegetation structure and function of alpine ecosystems are driven by regional and local contingencies in addition to macroclimatic factors.     

Hydraulic plasticity and limitations of alpine Rhododendron species

In the European Alps, Rhododendron ferrugineum grows in silicate regions while Rhododendron hirsutum is restricted to limestone areas. At geologically mixed sites, also hybrids (Rhododendron × intermedium) can occur. We hypothesised that hydraulic properties would vary with the species’ habitat requirements. Key hydraulic parameters (vulnerability to drought-induced embolism, stomata regulation) and related wood characteristics as well as diurnal courses of water potential (Ψ) and stomatal conductance were analysed on plants growing on a silicate, a limestone and a geologically mixed site. Highest embolism resistance[Ψ at 50% loss of conductivity (Ψ ₅₀), −3.24 ± 0.18 MPa] and the highest safety margin between the Ψ at stomata closure (Ψ _SC at 10% of maximal leaf conductance) and Ψ ₅₀ were observed in R. hirsutum at the limestone site (1.57 MPa). Like in R. ferrugineum, hydraulic parameters indicated less resistance at the geologically mixed site. Highest Ψ ₅₀ (−1.95 ± 0.12 MPa), corresponding to wide conduits and a reduced conduit wall reinforcement, was found in R. × intermedium. Diurnal courses indicated a rapid stomata closure in response to low Ψ in R. hirsutum and R. × intermedium. The plasticity in drought adaptation of R. hirsutum corresponds to its ability to colonise dry limestone areas. In contrast, hydraulic limitations of R. × intermedium may explain restrictions to rather moist sites. This study provides insight into the role of xylem hydraulics and stomata regulation in shrub water relations, interspecific and site-specific differences in drought adaptation, as well as effects of hybridisation on plant hydraulics.     

Nitrogen and carbon storage in alpine plants

Alpine plants offer unique opportunities to study the processesand economics of nutrient storage. The short alpine growingseason forces rapid completion of plant growth cycles, whichin turn causes competition between vegetative and reproductivegrowth sinks during the early part of the growing season. Mobilizationof stored nitrogen and carbon reserves facilitates competingsinks and permits successful completion of reproduction beforethe onset of winter stress. We discuss the theoretical frameworkfor assessing the costs and benefits of nutrient storage inalpine plants in order to lay the foundation for interpretationof observations. A principal point that has emerged from pasttheoretical treatments is the distinction between reserve storage,defined as storage that occurs with a cost to growth, and resourceaccumulation, defined as storage that occurs when resource supplyexceeds demand, and thus when there is no cost to growth. Wethen discuss two case studies, one already published and onenot yet published, pertaining to the storage and utilizationof nitrogen and carbon compounds in alpine plants from NiwotRidge, Colorado. In the first case, we tested the hypothesisthat the seasonal accumulation of amino acids in the rhizomeof N-fertilized plants of Bistorta bistortoides provides anadvantage to the plant by not imposing a cost to growth at thetime of accumulation, but providing a benefit to growth whenthe accumulated N is remobilized. We show that, as predicted,there is no cost during N accumulation but, not as predicted,there is no benefit to future growth. In the presence of N accumulation,reliance on stored N for growth increases, but reliance on current-season,soil-derived N decreases; thus the utilization of availableN in this species is a ‘zero sum’ process. Inherentmeristematic constraints to growth cause negative feedback thatlimits the utilization of accumulated N and precludes long-termadvantages to this form of storage. In the second case study,we discuss new results showing high concentrations of cyclicpolyol (cyclitol) compounds in the leaves of many alpine speciesdominant in the dry fellfield habitat. In Artemisia scopulorum,cyclitols were induced as the growing season progressed, andreached highest concentrations during the dry, late-summer months.Leaf cyclitol concentrations were high in all four species ofthe Caryophyllaceae that we examined and appeared to be constitutivecomponents of the leaf carbohydrate pool as concentrations werehigh through the entire growing season. We observed correlationsamong seedling abundance, seeding survivorship and the presenceof high leaf cyclitol concentrations. We propose that the primaryfunction of cyclitols in the leaves of alpine, fellfield herbsis to promote drought tolerance through osmotic protection,and enhance fitness by improving seedling survival. We consideredthe possibility that cyclitols also function as carbon storagecompounds that are remobilized at the end of the growing seasonand used to support growth the following year. Our observationsdo not support this hypothesis in the Caryophyllaceae becausethe requirement for high constitutive concentrations year-after-yearprevents long-term advantages of storage and remobilization.However, in A. scopulorum, remobilization of cyclitols followingthe end of the growing season may provide storage substratesthat can be used for growth the following season. From our analysiswe conclude that it is difficult to use current theory thatis embedded in the economic concept of costs and benefits tointerpret observed dynamics in nitrogen and carbon allocation.Future theoretical developments that move away from an abstractfoundation embedded in cost-benefit tradeoffs and toward phenotypicintegration of source-sink relationships will improve our abilityto merge observations and theory.