ENVIRONMENTAL AND PHYSIOLOGICAL FACTORS INFLUENCING THE NATURAL DISTRIBUTION OF EVERGREEN AND DECIDUOUS ERICACEOUS SHRUBS ON NORTHEAST AND SOUTHWEST SLOPES OF THE SOUTHERN APPALACHIAN MOUNTAINS. I. IRRADIANCE TOLERANCE

Deciduous and evergreen species are segregated on northeast and southwest slopes of the southern Appalachian Mountains. The segregated distributions of three ericaceous shrubs (Rhododendron maximum valley positions; Rhododendron periclymenoides on northeast slopes; Kalmia latifolia on southwest slopes) were compared to the respective irradiance environments. Growth patterns of field plants, and photosynthetic acclimation of each species to three irradiance treatments in a phytotron were studied. Rhododendron maximum, an evergreen species, was found to be most sensitive to high radiation. In phytotron experiments, quantum yield, light saturated photosynthetic capacity, photosynthesis per chlorophyll, and water use efficiency decreased at high ambient irradiance for R. maximum. These characteristics limit the growth of R. maximum on high irradiance southwestern slopes. Both K. latifolia and R. periclymenoides were able to improve their photosynthetic performance at high ambient irradiance. Rhododendron periclymenoides, a deciduous species, was found to continue increasing leaf conductance at high irradiance without an increase in photosynthesis indicating a possible limitation by water in high light environments such as southwest slopes. Kalmia latifolia, an evergreen species, had reduced photosynthetic capacity and reduced water use efficiency when grown in low irradiance conditions which coincides with the higher K. latifolia abundance on high light, southwestern slopes.     

INFLUENCE OF AGE AND MICROCLIMATE ON THE PHOTOCHEMISTRY OF RHODODENDRON MAXIMUM LEAVES II. CHLOROPLAST STRUCTURE AND PHOTOSYNTHETIC LIGHT RESPONSE

The influence of age on chloroplast structure and photosynthetic light response of Rhododendron maximum L. was studied in three different microhabitats. The three microhabitats constituted a gradient of low, intermediate, and high irradiance levels. The most dramatic change in chloroplast structure with increasing age was the proliferation of the number and size of plastoglobuli. The magnitude and age specific rate of chloroplast occlusion by plastoglobuli increased in habitats with higher irradiance. Photosynthetic responses to light differed among the age categories of leaves. Light saturated photosynthesis and quantum yield decreased as leaves aged. However, in high light environments the rate of reduction of quantum yield or light saturated photosynthetic rate was more rapid than in the low light environment. The quantity of plastoglobuli increased in association with reduced light reaction capacity. The presence and abundance of plastoglobuli in R. maximum chloroplasts and their association with reduced photosynthetic performance indicates that the photosynthetic apparatus of the R. maximum chloroplast is sensitive to photodestruction by high irradiance: commonly a winter phenomenon in these environments.     

Seasonal and diurnal leaf movements of Rhododendron maximum L. in contrasting irradiance environments

Summary Leaf orientation (azimuth and angle) and leaf curling were measured seasonally and diurnally on Rhododendron maximum L. under an evergreen and a deciduous canopy. The microclimatic conditions under the evergreen canopy (mixed pine and hemlock) were characterized by lower irradiance but similar temperature, and vapor pressure deficit (vpd) to that under the deciduous canopy (mixed oak and maple). Under both canopies irradiance was more intense during winter months.On a seasonal basis leaf angle was closer to horizontal under the evergreen canopy but there was no difference between leaf curling in the two sites. Stomatal conductance was higher under the deciduous canopy but stomata were closed in the winter (following canopy abscission) under the evergreen and deciduous canopies even during warm winter days. Leaf water potentials were lower in the winter and Rhododendron maximum had higher leaf water potentials under the evergreen canopy.Significant association between mean leaf angle and curling index were found above a mean leaf angle of 70°. Leaf curling was highly associated with leaf temperature where 0° C was a critical value stimulating leaf curling. Leaf angle was linearly related to leaf temperatures above 0° C although this relationship was different under the two canopy types as a result of differing irradiance or differing water potential.     

QUANTITATIVE PHENOLOGY AND LEAF SURVIVORSHIP OF RHODODENDRON MAXIMUM IN CONTRASTING IRRADIANCE ENVIRONMENTS OF THE SOUTHERN APPALACHIAN MOUNTAINS

Seasonal plant growth patterns were compared for Rhododendron maximum L. in two contrasting subcanopy environments. The two subcanopy, above ground environments differed only in their quantity of irradiance by virtue of the relative dominance of evergreen or deciduous trees in the canopy. A third site had no canopy influence. Overall growth (shoot elongation, woody increment, leaf production) was maximized under the open (BMO) and deciduous dominated canopy (PCD). The leaf pool was significantly smaller under the evergreen dominated canopy (PCE) but average leaf area per leaf was slightly larger at PCE. Individual age-specific leaf cohorts, identified from shoot morphology, indicated increased leaf survivorship with a decreased irradiance environment. Leaf production was synchronous and rapid (1 week), followed by three weeks of leaf expansion, which created the even-aged leaf cohorts. Wood growth (diameter increment), in contrast, continued through the beginning of the winter. Reproductive effort increased with increasing irradiance environment. Significant variation in growth was observed between canopy shoot types at all three research sites. The significance of these phenological patterns is discussed in view of the variable subcanopy environment of southwestern Virginia.     

Leaf phenology and seasonal variation of photosynthesis of invasive <Emphasis Type="Italic">Berberis thunbergii</Emphasis> (Japanese barberry) and two co-occurring native understory shrubs in a northeastern United States deciduous forest

Early leafing and extended leaf longevity can be important mechanisms for the invasion of the forest understory. We compared the leaf phenology and photosynthetic characteristics of Berberis thunbergii, an early leafing invasive shrub, and two co-occurring native species, evergreen Kalmia latifolia and late leafing Vaccinium corymbosum, throughout the 2004 growing season. Berberis thunbergii leafed out 1 month earlier than V. corymbosum and approximately 2 weeks prior to the overstory trees. The photosynthetic capacity [characterized by the maximum carboxylation rate of Rubisco (V _cmax) and the RuBP regeneration capacity mediated by the maximum electron transport rate (J _max)] of B. thunbergii was highest in the spring open canopy, and declined with canopy closure. The 2003 overwintering leaves of K. latifolia displayed high V _cmax and J _max in spring 2004. In new leaves of K. latifolia produced in 2004, the photosynthetic capacity gradually increased to a peak in mid-September, and reduced in late November. V. corymbosum, by contrast, maintained low V _cmax and J _max throughout the growing season. In B. thunbergii, light acclimation was mediated by adjustment in both leaf mass per unit area and leaf N on a mass basis, but this adjustment was weaker or absent in K. latifolia and V. corymbosum. These results indicated that B. thunbergii utilized high irradiance in the spring while K. latifolia took advantage of high irradiance in the fall and the following spring. By contrast, V. corymbosum generally did not experience a high irradiance environment and was adapted to the low irradiance understory. The apparent success of B. thunbergii therefore, appeared related to a high spring C subsidy and subsequent acclimation to varying irradiance through active N reallocation and leaf morphological modifications.     

Comparative field study of spring and summer leaf gas exchange and photobiology of the Mediterranean trees Quercus ilex and Phyllyrea latifolia

Summer-induced changes in gas exchange, fluorescence and reflectance were measured on leaves of two co-occurring Mediterranean small trees, Quercus ilex and Phillyrea latifolia, in May, June and July 1996 in Central Catalonia (NE Spain). The humid 1996 summer only produced mild water stress conditions. However, photosynthesis (A) and stomatal conductance (gs) decreased in June and July in both species. In June P. latifolia had higher net photosynthetic rates and lower stomatal conductances than Q. ilex, thus exhibiting higher instantaneous plant water use efficiencies. In agreement with these results, the photochemical reflectance index (PRI, calculated as (R570-R531)/(R531+R570)) of P. latifolia was lower, suggesting a possible lower xanthophyll de-epoxidation state. However, P. latifolia had lower F/F'm and therefore a lower electron transport rate (ETR). The behaviour of PRI confirmed previous studies indicating a strong relationship between PRI, F/F'm and photosynthetic radiation-use efficiency (PRUE). PRI offers a simple, portable means of assessing PRUE with the potential for remote sensing applications. Finally, the possible ecological consequences of these results on the behaviour of the two species studied under the predicted warmer and drier conditions of global change are discussed.Key words: Quercus ilex, Phillyrea latifolia, summer, net photosynthetic rates, stomatal conductance, intrinsic water use efficiency, photosynthetic radiation-use-efficiency, reflectance, fluorescence, photochemical reflectance index, photochemical efficiency, water index.      

Coordination of leaf N,anatomy, photosynthetic capacity,and hydraulics enhances evergreen expansive potential

Key message

Reduced leaf longevity, N-fixation, and enhanced hydraulic capacity combined support greater shifts in seasonal photosynthetic capacity of an expansive understory evergreen woody species relative to co-occurring less expansive evergreen species.

Abstract

Physiological functioning typically declines with increased leaf life span. While an evergreen leaf habit is generally associated with reduced leaf N, physiological capacity, and slower growth, most expansive woody species are evergreens and/or N fixers. An evergreen leaf habit enables year-round activity and less investment in carbon and nutrients, while N-fixation enhances photosynthetic capacity. Our objective was to compare anatomy and physiology of three woody evergreens Ilex opaca Aiton (Aquifoliaceae), Kalmia latifolia L. (Ericaceae), and Myrica cerifera (Myricaceae) of varying leaf longevity, N-fixation capability, and known expansive potential in a deciduous forest understory to determine if seasonal physiological performance integrated these factors. We hypothesized that I. opaca (non-expansive) and K. latifolia (moderately expansive), which have longer leaf longevities, would have reduced physiological performance compared to M. cerifera (expansive), which has shorter leaf longevity, and symbiotically fixes atmospheric N. Stomatal conductance to water vapor, photosynthetic and hydraulic capacities, specific leaf area, and leaf %N decreased with increasing leaf life span; however, trends among species were not consistent seasonally. While hydraulic capacity remained constant throughout the year, photosynthetic capacity did not. During the growing season, M. cerifera displayed photosynthetic capacity similar to deciduous species, yet, during the winter, photosynthetic capacity was similar to the slower-growing evergreens. Reduced leaf life span, enhanced hydraulic capacity, and nitrogen fixation support the seasonal shift in photosynthetic capacity observed in M. cerifera. This “hybrid” strategy enables M. cerifera to maximize productivity during months of optimal conditions, thereby promoting rapid growth and expansion in the understory.     

环境因子对小麦叶片水分利用效率影响的实验研究和数值模拟

根据现有的光合作用和蒸腾作用的模型,利用Ｂａｌｌ－Ｂｅｒｒｙ 的气孔导度模型,将叶片的光合作用模型和蒸腾作用模型结合起来,建立了光强（Ｉ）、叶片－大气水汽饱和差（ＶＰＤ）和大气ＣＯ２ 浓度（Ｃａ）等环境因子对小麦叶片水分利用效率（ＷＵＥ）影响的模型。由于这３ 种环境因子对光合、蒸腾的影响方式上的差异,作为两者之比的叶片ＷＵＥ随各环境因子的变化出现复杂的图景,同时,在人工气候箱内分别于这３ 个因子变化时对光合、蒸腾的变化作了测定,计算了叶片的Ｗ ＵＥ。测定结果与模拟结果的对比表明,在多数情况下两者符合程度良好,但在高Ｃａ下有较大的偏离     

DIFFERENCES IN LEAF STRUCTURE,CHLOROPHYLL, AND NUTRIENTS FOR THE UNDERSTORY TREE ASIMINA TRILOBA

To investigate differences in leaf structure, chlorophyll and nutrients on terminal branches of the understory tree Asimina triloba, the first (proximal) and the last (distal) leaves to develop in the spring were compared. Proximal leaf expansion was completed before the overstory canopy was fully closed but distal leaf expansion occurred during and after the development of the overstory canopy. Fully expanded proximal leaves were 76% smaller in area, were 18% thicker and had 36% more stomates per m of leaf area when compared to distal leaves. In addition, maximum stomatal conductance to water vapor was greater (150 vs. 120 mmol m^−-2s^−-1) and the minimum PPFD required for maximum conductance was higher (200 vs. 150 μmol m^−-2s^−-1) for the proximal leaves. Chlorophyll content was also greater for proximal leaves, but nitrogen and phosphorus contents were lower throughout the entire summer. Seasonal measurements indicated an increase in chlorophyll a content and reductions in nitrogen content throughout the summer growth period for leaves from both positions. The results suggest that distal and proximal leaves differed physiologically and that the measured differences were related to the changing irradiance environment during leaf development. The time of leaf expansion, as indicated by leaf position on the branch, may be an important consideration when examining the water and photosynthetic relations of understory trees.     

Growth and Leaf Gas Exchange Characteristics in Dalbergia sissoo Roxb. and D. latifolia Roxb. Under Water Deficit

Forty two-month-old plants of Dalbergia sissoo and D. latifolia were subjected for 56 d to water deficit induced by withholding water. Drought stress caused a significant reduction in plant height, stem diameter, net photosynthetic rate (P _N), transpiration rate (E), and stomatal conductance (g _s) in both species, but the reduction was greater in D. sissoo than in D. latifolia. Water use efficiency (P _N/E) was adversely affected due to water stress only in D. latifolia, and intrinsic water use efficiency (P _N/g _s) was increased in both species. There was a slight effect of water stress on variable to maximum fluorescence (F_v/F_m) (quantum yield of photosystem 2) in both species, but the species did not differ significantly in this attribute.     

Arundo donax L. (Poaceae) — a C3 Species with Unusually High Photosynthetic Capacity

Gas exchange characteristics, chlorophyll a fluorescence and leaf water potential were investigated in the giant reed, Arundo donax, under natural conditions in an estuarine mangrove swamp in Durban, South Africa. Maximum photosynthetic CO₂ uptake ranged between 19.8 and 36.7 μmol m^?2 s^?1, depending on irradiance, and appeared to be regulated by leaf conductance. There was no saturation of CO₂ uptake or electron transport through PSII (ETR) with increasing irradiance up to 2500 μmol photons m^?2 s^?1. A linear relationship between CO₂ uptake, corrected for respiration (A), and ETR has only been reported for C₄ species and C₃ species when photorespiration is eliminated. From this relationship, it was calculated that 8.5 electrons were transported through PSII for the fixation of one mole of CO₂. Predawn leaf water potential was about ?0.5 MPa and decreased to ?1.5 MPa on a cloudy day and to ?2.1 MPa on a clear day. Diurnal change in leaf water potential had little influence on leaf conductance and hence CO₂ uptake. The molar water use efficiency (WUE) ranged between 4.1 and 9.3 μmol mmol^?1. Percentage photorespiration was between 36 and 39%.     

Influence of water relations and temperature on leaf movements of rhododendron species

Rhododendron maximum L. and R. Catawbiense L. are subcanopy evergreen shrubs of the eastern United States deciduous forest. Field measurements of climate factors and leaf movements of these species indicated a high correlation between leaf temperature and leaf curling; and between leaf water potential and leaf angle. Laboratory experiments were performed to isolate the influence of temperature and cellular water relations on leaf movements. Significant differences were found between the patterns of temperature induction of leaf curling in the two species. Leaves of the species which curled at higher temperatures (R. catawbiense) also froze at higher leaf temperatures. However, in both cases leaf curling occurred at leaf temperatures two to three degrees above the leaf freezing point. Pressure volume curves indicated that cellular turgor loss was associated with a maximum of 45% curling while 100% or more curling occurred in field leaves which still had positive cell turgor. Moisture release curves indicated that 70% curling requires a loss of greater than 60% of symplastic water which corresponds to leaf water potentials far below those experienced in field situations. Conversely, most laboratory induced changes in leaf angle could be related to leaf cell turgor loss.     

A Comparison of Photosynthetic Capacity between Myrica Gale var. Tomentosa a Nitrogen-Fixing Shrub,and Co-Occurring Species in an Oligotrophic Moor

Photosynthetic and transpiration (E) rates, stomatal conductance, and leaf nitrogen content were surveyed for Myrica gale var. tomentosa, a N₂-fixing wetland shrub, Betula platyphylla var. japonica, and Rhododendron japonicum in Ozegahara moor, an oligotrophic moor in Central Japan. Net photosynthetic rate saturated with irradiance (P_max) of M. gale was 15.2-16.5 μmol(CO₂) m^-2 s^-1, higher than those of the other species throughout the growing season. P_max was positively correlated with leaf N content among the three species. The large leaf N content in M. gale was due to N₂-fixation in root nodules. In a comparison of M. gale in two habitats, P_max, leaf N content, and root nodule development were larger in the wetter habitat. M. gale showed high E and no midday depression of P_max even under high irradiance and large vapour pressure deficit between leaves and ambient air on a midsummer day. These traits of photosynthesis and water relations were associated with the dominance of this shrub in wetter sites such as stream sides and hollows. This revised version was published online in September 2006 with corrections to the Cover Date.     

不同光照条件下喜阴植物谢君魔芋对稳态和动态光源的响应特征

谢君魔芋(Amorphophallus xiei)是起源于云南西南地区热带雨林的典型喜阴植物,近年来得到了广泛种植和推广,在种植过程中,谢君魔芋需要采用遮荫栽培模式。为了揭示谢君魔芋对光照强度的适应策略,该研究探讨了生长在不同光照强度下(透光率为50%、29%、17%、7%)谢君魔芋叶片的光合作用特征、光合诱导特征、光合色素含量以及叶片氮素(N)含量和N分配。结果表明:随着生长环境光照强度的降低,单位叶面积和单位叶质量最大净光合速率、光合色素含量、最大羧化速率、最大电子传递速率及比叶面积均增大,而暗呼吸和光补偿点均减小。在光合诱导过程中,生长在透光率为17%光环境中的谢君魔芋完成50%光合诱导所需的时间最短,约为81.4 s;在光诱导进行10 min时,诱导状态最高,为87.3%。完成50%和90%光合诱导所需的时间与低光下初始气孔导度呈负相关关系。随着生长光照强度降低,叶片中的N分配到羧化组分和生物能转化组分中的比例先增大后减小,在透光率为17%的光环境下具有最大值;而叶片中的N分配到捕光色素组分中的比例随着生长环境光照强度降低而增加。该研究结果表明,喜阴植物谢君魔芋通过加强对低光和动态光源的利用能力及有效的N资源分配策略来适应低光照环境。     

Carbon limitation leads to suppression of first year oak seedlings beneath evergreen understory shrubs in Southern Appalachian hardwood forests

Inhibition of canopy tree recruitment beneath thickets of the evergreen shrubs Rhododendron maximum L. and Kalmia latifolia L. has long been observed in Southern Appalachian forests, yet the mechanisms of this process remain unresolved. We present a first-year account of suppression of oak seedlings in relation to Rhododendron and Kalmia basal area, light and resource availability, seedling performance and the rates of seedling damage (i.e., herbivory). We found no evidence of first-year seedling suppression or significant resource deficiencies beneath thickets of K. latifolia in mature mixed hardwood stands. Suppression beneath R. maximum was apparent during the first growing season. We found that seedling biomass, light availability prior to canopy closure, and seedling tissue C:N ratios were negatively correlated with R. maximum basal area. Basal area of R. maximum was positively correlated with seedling mortality rates, soil [Al], and early-growing season leaf herbivory rates. Seedling growth was positively correlated with light and tissue C:N, while negatively correlated with soil [Al]. Overall, our results support the inhibition model of shade-mediated carbon limitation beneath dense understory shrubs and indicate the potential importance of herbivory and aluminum toxicity as components of a suppression mechanism beneath R. maximum thickets. We present a causal model of first year inhibition beneath R. maximum in the context of our findings and the results of prior studies.     

EFFECTS OF CANOPY POSITION AND IRRADIANCE ON THE LEAF PHYSIOLOGY AND MORPHOLOGY OF PENTACLETHRA MACROLOBA (MIMOSACEAE)

Pentaclethra macroloba (Willd.) Kuntze (Mimosaceae) is a dominant late-successional tree species in the Atlantic lowland forests of Costa Rica. Leaves of P. macroloba from three heights in the forest canopy were compared with leaves of seedlings grown in controlled environment chambers under four different irradiance levels. Changes in leaf characteristics along the canopy gradient paralleled changes resulting from the light gradient under controlled conditions. The effect of light or canopy position on light-saturated photosynthesis was small, with maximum photosynthesis increasing from 5 to 6.5 μmol m^−-2 s^−-1 from understory to canopy. Both chamber grown and field leaves showed large adjustments in photosynthetic efficiency at low light via reductions in dark respiration rates and increases in apparent quantum yields. Light saturation of all leaves occurred at or below 500 μmol m^−-2 s^−-1. Leaf thickness, specific leaf weight, and stomatal density increased to a greater extent than saturated photosynthesis with higher irradiance during growth or height in the canopy. As a result, there was a poor correspondence between leaf thickness and light-saturated photosynthesis on an area basis. It is concluded that Pentaclethra macroloba possesses the characteristics of a typical shade-tolerant species.     

Photosynthetic limitation of several representative subalpine species in the Catalan Pyrenees in summer

Information on the photosynthetic process and its limitations is essential in order to predict both the capacity of species to adapt to conditions associated with climate change and the likely changes in plant communities. Considering that high‐mountain species are especially sensitive, three species representative of subalpine forests of the Central Catalan Pyrenees: mountain pine (Pinus uncinata Mill.), birch (Betula pendula Roth) and rhododendron (Rhododendron ferrugineum L.) were studied under conditions associated with climate change, such as low precipitation, elevated atmospheric [CO₂] and high solar irradiation incident at Earth's surface, in order to detect any photosynthetic limitations. Short‐term high [CO₂] increased photosynthesis rates (A) and water use efficiency (WUE), especially in birch and mountain pine, whereas stomatal conductance (g_s) was not altered in either species. Birch showed photosynthesis limitation through stomatal closure related to low rainfall, which induced photoinhibition and early foliar senescence. Rhododendron was especially affected by high irradiance, showing early photosynthetic saturation in low light, highest chlorophyll content, lowest gas exchange rates and least photoprotection. Mountain pine had the highest A, photosynthetic capacity (A_max) and light‐saturated rates of net CO₂ assimilation (A_sat), which were maintained under reduced precipitation. Furthermore, maximum quantum yield (F_v/F_m), thermal energy dissipation, PRI and SIPI radiometric index, and ascorbate content indicated improved photoprotection with respect to the other two species. However, maximum velocity of carboxylation of RuBisco (V_cmax) indicated that N availability would be the main photosynthetic limitation in this species.     

The relationship between freezing tolerance and thermotropic leaf movement in five Rhododendron species

Summary Leaf movement kinetics in five species of Rhododendron were studied in response to leaf temperature, leaf freezing point, and leaf water deficit. There was a gradient in the degree of leaf curling among species in the following order from the greatest curling to the least curling: Rhododendron catawbiense, R. maximum, R. minus, R. macrophyllum, R. ponticum. Those species found to be tolerant of winter conditions had the most intense leaf movements (both curling and angle) while those species with minimal cold tolerance had limited or no leaf movements. Leaf curling occurred at leaf temperatures above the tissue freezing points in all species. Athough leaf angle was influenced by leaf turgor, general tissue desiccation was not the ultimate cause for thermotropic leaf curling in any species tested. Those species with the greatest leaf curling and angle movements had the highest osmotic potential, the lowest water deficit at the turgor loss point, and the lowest symplastic water fraction. These data suggest that there is a trade off in Rhododendron leaf physiology between cold tolerance (due to leaf movements) and water stress tolerance (due to turgor maintenance mechanisms).     

Establishment of co‐existing Mediterranean tree species under a varying soil moisture regime

We investigated the responses of two co‐existing Mediterranean trees with different regeneration strategies (Phillyrea latifolia seedlings and Quercus ilex sprouts) to experimental drought below the forest canopy. We considered different recruitment stages and used leaf isotopic discrimination to estimate water use efficiency (WUE) and nitrogen availability and use. Drought decreased the emergence and survival of seedlings and sprouts. Survival and growth of older saplings were not influenced by drought. Seedling emergence of P. latifolia was higher than Q. ilex sprout production, but Q. ilex sprouts had higher survival and growth rates. These differences disappeared in the sapling stage. Carbon isotopic discrimination suggested that Q. ilex sprouts had higher WUE than P. latifolia seedlings. Drought increased WUE of recruits, particularly in Q. ilex. Water use regulation increased with ageing, particularly in P. latifolia. Q. ilex had higher δ¹⁵N values than P. latifolia; these were also higher under drier soil conditions. Current year seedlings had higher δ¹⁵N than saplings, particularly in P. latifolia, suggesting they exploit superficial soil layers. These results suggest that sprouts obtain benefit from resources stored in parent plants. At earlier stages, they perform better than seedlings. This response is not coupled to adult vulnerability to drought for these species, revealing the difficulty of predicting species dynamics during climate change.     

Leaf traits of natural populations of <Emphasis Type="Italic">Adiantum reniforme</Emphasis> var. <Emphasis Type="Italic">sinensis</Emphasis>, endemic to the Three Gorges region in China

Leaf mass per unit area (LMA), carbon and nitrogen contents, leaf construction cost, and photosynthetic capacity (P _max) of Adiantum reniforme var. sinensis, an endangered fern endemic to the Three Gorges region in southwest China, were compared in five populations differing in habitat such as soil moisture and irradiance. The low soil moisture and high irradiance habitat population exhibited significantly higher LMA, area-based leaf construction (CC_A), and carbon content (C_A), but lower leaf nitrogen content per unit dry mass (N_M) than the other habitat populations. The high soil moisture and low irradiance habitat populations had the lowest CC_A, but their cost/benefic ratios of CCA/P _max were similar to the medium soil moisture and irradiance habitat population due to their lower leaf P _max. Hence A. reniforme var. sinensis prefers partially shaded, moist but well-drained, slope habitats. Due to human activities, however, its main habitats now are cliffs or steeply sloped bare rocks with poor and thin soil. The relatively high energy requirements and low photosynthetic capacity in these habitats could limit the capability of the species in extending population or interspecific competition and hence increase its endangerment.