MICROCLIMATIC EFFECTS ON WATER RELATIONS,LEAF TEMPERATURES,AND THE DISTRIBUTION OF HERACLEUM LANATUM AT HIGH ELEVATIONS

The small-scale distribution of an understory herb, Heracleum lanatum, was evaluated in terms of leaf temperature and water relations limitations due to a large leaf size (630 cm²). Diurnal variations in transpiration (4 to 60 mg m⁻² s⁻¹) were influenced by fluctuations in solar irradiance, wind speed, leaf temperature and stomatal conductance. Computer simulations indicated that leaf temperatures in a forest clearing would be > 12 C above air temperature, with maximum transpiration rates of 140 mg m⁻² s⁻¹, and daily water loss to be over 200% greater than values at natural understory locations. Simulations of nocturnal temperature relations indicated ~100 W m ⁻² less incident longwave irradiance in the forest clearing as compared to the understory (560 vs. 660 W m⁻² at 400 hr). This difference led to predicted leaf temperatures being as low as 6 C below air temperature in the forest clearing while measured leaf temperatures in the forest understory were within 1.5 C of air temperature throughout the night. Furthermore, minimum air temperatures were at or below 6 C on 36% of the nights during the summer growth period indicating that in open areas leaves of H. lanatum would frequently be below 0 C and subject to possible freeze damage. Heracleum lanatum may be more abundant in the shaded understory of the subalpine forest because exposure in open environments would result in high leaf temperatures and increased transpirational water loss during the day, as well as low leaf temperatures with the possibility of freeze damage at night.     

Does Prunus serotina act as an aggressive invader in areas with a low propagule pressure?

Since most studies on Prunus serotina in Western Europe focused on heavily invaded areas, we wondered whether P. serotina also acts as an aggressive invader in areas with a low propagule pressure. Based on long-term data for the Liedekerke forest reserve, we found that connectivity to seed sources and light availability were the major drivers of P. serotina presence: long-distance dispersal events and ‘windows of opportunity’ seem to direct P. serotina colonization. In the studied forest, P. serotina could not be considered an aggressive invader since its spread slowed down rather quickly and did not hamper the establishment of native tree species. Furthermore, understory P. serotina showed low growth and seed production, while the high Rubus cover hampered germination and establishment. Nonetheless, calamities opening up the canopy layer in the few areas with high P. serotina sapling density might alter the course of the invasion process.     

Stomatal acclimation to vapour pressure deficit doubles transpiration of small tree seedlings with warming

Future climate change is expected to increase temperature (T) and atmospheric vapour pressure deficit (VPD) in many regions, but the effect of persistent warming on plant stomatal behaviour is highly uncertain. We investigated the effect of experimental warming of 1.9–5.1 °C and increased VPD of 0.5–1.3 kPa on transpiration and stomatal conductance (g_s) of tree seedlings in the temperate forest understory (Duke Forest, North Carolina, USA). We observed peaked responses of transpiration to VPD in all seedlings, and the optimum VPD for transpiration (D_opt) shifted proportionally with increasing chamber VPD. Warming increased mean water use of Carya by 140% and Quercus by 150%, but had no significant effect on water use of Acer. Increased water use of ring‐porous species was attributed to (1) higher air T and (2) stomatal acclimation to VPD resulting in higher g_s and more sensitive stomata, and thereby less efficient water use. Stomatal acclimation maintained homeostasis of leaf T and carbon gain despite increased VPD, revealing that short‐term stomatal responses to VPD may not be representative of long‐term exposure. Acclimation responses differ from expectations of decreasing g_s with increasing VPD and may necessitate revision of current models based on this assumption.     

Variations in leaf morpho‐anatomy and photosynthetic traits between sun and shade populations of Eurya japonica (Pentaphylacaceae) whose seeds are dispersed by birds across habitats

Eurya japonica occurs in diverse light environments through seed dispersal by birds. As the seed size is extremely small, we hypothesized that newly germinated seedlings with restricted depth of roots and length of the hypocotyl would suffer high mortality due to increased transpiration in sunny habitats and low light in shady habitats. We also expected that surviving seedlings would differ in leaf traits between habitats as a result of selection. We aimed to determine how photosynthetic traits differ between habitats and how leaf structure is related to this difference. We examined photosynthesis and leaf morpho‐anatomy for plants cloned from cuttings collected from the forest understory (shade population) and neighboring roadsides and cut‐over areas (sun population) and then grown under two irradiances (18.5% and 100% sunlight) in an experimental garden. Under growth in 100% sunlight, cloned plants from the sun population exhibited significantly greater area‐based photosynthetic capacity compared to cloned plants from the shade population at a comparable stomatal conductance, which was attributable to a higher area‐based leaf nitrogen concentration. On the other hand, mean values of photosynthetic capacity did not significantly differ between the two populations. Cloned plants from the sun population had significantly thicker leaf laminas and spongy tissue and lower stomatal density compared to cloned plants from the shade population. Thickened leaf lamina might have increased leaf tolerance to physical stresses in open habitats. The variation in leaf morpho‐anatomy between the two populations can be explained in terms of the economy of leaf photosynthetic tissue.     

Photosynthetic acclimation to light in juveniles of two cloud forest tree species

The photosynthetic response of juveniles of Decussocarpus rospigliosii, an emergent primary forest species and shade tolerant in its juvenile stages and Alchornea triplinervia, a gap-colonizing species of tropical cloud forest in Venezuela was studied. Daily courses of microenvironmental variables and gas exchange under contrasting light conditions (gap and understory) were carried out in their natural environment and transplanted to different light regimes (shade and sun) in the field. The photosynthetic response and some anatomical characteristics of plants from different treatments were analyzed in the laboratory. Photosynthetic rates were low for both species, and were negative during some diurnal periods, related to the low photosynthetically active radiation levels obtained at both gap (6% of total radiation) and understory (2%). A. triplinervia shows higher rates (1.5–3.0 molm^{-2 -1}) than D. rospigliosii (0.7–1.1 molm^-2s^-1). Both species showed increased photosynthetic rates when grown in gaps. A. triplinervia did not adjust its maximum photosynthetic rates to the prevailing light conditions. In contrast, D. rospigliosii responded to increased light levels. Both species showed low light compensation points when grown under total shade. There was a partial stomatal closure generally during midday in D. rospigliosii. A. triplinervia presented lower leaf conductances, transpiration rates and lesser stomatal control. Some leaf anatomical characteristics, in both species, were affected by variations in the light regime (i.e. increased leaf thickness, leaf specific weight and stomatal density). These results suggest that both species have the ability to respond to variations in their natural light environments, therefore maintaining a favorable carbon balance during the day.     

Losses of seeds of temperate trees to soil fungi: effects of habitat and host ecology

Evidence suggests that impacts of fungal pathogens on tree recruitment tend to be greater in the forest understory than in openings, and that shade-tolerant trees are less vulnerable than shade-intolerant species. To investigate the role that harmful soil fungi may have in reducing regeneration of temperate trees, we applied fungicide to buried seeds of matched pairs of species differing in their relative shade tolerance and/or successional status (Acer negundo versus Acer saccharum, Prunus virginiana versus Prunus serotina, and Pinus strobus versus Tsuga canadensis), in three habitats that differed in their degree of openness (old field, forest gap, and forest understory). Our results indicated that soil fungi reduced germination of A. negundo, P. virginiana, P. serotina. and T. canadensis, and reduced viability of ungerminated seeds of P. strobus; no significant effects of fungi on seeds of A. saccharum were detected. However, we found seeds were not less likely to survive following burial in forest understory than in gaps. As well, results for only one species pair (A. negundo versus A. saccharum) were consistent with the prediction that shade-intolerant or successional species should be more susceptible to fungal attack than mature forest species. These results contrast with other studies of temperate and especially tropical forest trees.     

Understory Invasion by <Emphasis Type="Italic">Acacia longifolia</Emphasis> Alters the Water Balance and Carbon Gain of a Mediterranean Pine Forest

In water-limited ecosystems, where potential evapotranspiration exceeds precipitation, it is often assumed that plant invasions will not increase total ecosystem water use, because all available water is evaporated or transpired regardless of vegetation type. However, invasion by exotic species, with high water use rates, may potentially alter ecosystem water balance by reducing water available to native species, which may in turn impact carbon assimilation and productivity of co-occurring species. Here, we document the impact of invasion by an understory exotic woody species (Acacia longifolia) in a semi-arid Mediterranean dune pine forest. To quantify the effects of this understory leguminous tree on the water use and carbon fixation rates of Pinus pinaster we compare an invaded and a non-invaded stand. A. longifolia significantly altered forest structure by increasing plant density and leaf area index in the mid-stratum of the invaded forest. A. longifolia contributed significantly to transpiration in the invaded forest (up to 42%) resulting in a slight increase in stand transpiration in the invaded relative to non-invaded forest. More importantly, both water use and carbon assimilation rates of P. pinaster were significantly reduced in the invaded relative to non-invaded stand. Therefore, this study shows that exotic plant invasions can have significant impacts on hydrological and carbon cycling even in water-limited semi-arid ecosystems through a repartitioning of water resources between the native and the invasive species.     

Vegetation effects on microclimate in lowland tropical forest in Costa Rica

The temperature and atmospheric humidity in a tropical lowland rain forest in Costa Rica were measured in order to assess the microclimate in different forest environments. Two disturbed sites, a single tree fall gap (400 m²) and an 0.5 hectare clearing, were compared for periods up to two years after disturbance. Two locations in primary forest, the canopy and understory, were also monitored. Temperatures were highest in the clearing, intermediate in the canopy and gap which were similar, and lowest in the understory. Vapor pressure deficits (VPD) were highest in the clearing, followed by the canopy, the gap and the understory. With regrowth of the vegetation in the gap and clearing sites, the temperatures and vapor pressure deficits significantly decreased. After 1 year, the microclimate at seedling height in the clearing resembled that of the gap, and after two years the microclimate of the gap was very similar to that of the understory. Seasonal differences in temperature and VPD were small compared to differences caused by changes in the stature of the vegetation.     

Is distribution of hydraulic constraints within tree crowns reflected in photosynthetic water-use efficiency? An example of <Emphasis Type="Italic">Betula pendula</Emphasis>

Spatial and daily variation in photosynthetic water-use efficiency was examined in leaves of Betula pendula Roth with respect to distribution of hydraulic conductance within the crown, morphological properties of stomata, and water availability. Intrinsic water-use efficiency (A _n/g _s) was determined from gas-exchange measurements performed both in situ in a natural forest stand and on detached shoots under laboratory conditions. In intact foliage, sun leaves demonstrated significantly higher (P < 0.001) A _n/g _s than shade leaves, as photosynthesis in the lower canopy was chronically limited by low light availability. However, this difference reversed in the mid-day period under sufficient irradiance (I > 800 μmol m⁻² s⁻¹): A _n/g _s averaged 28.8 and 24.0 μmol mol⁻¹ (P < 0.01) for shade and sun leaves, respectively. This last finding coincided with the data obtained in laboratory conditions: under equivalent leaf water supply and light, A _n/g _s in shade foliage was greater (P < 0.001) than in sun foliage across a wide range of irradiance. Thus, shade foliage of B. pendula is characterized by inherently higher A _n/g _s than sun foliage, associated with more conservative stomatal behavior, and lower soil-to-leaf (K _T) and leaf hydraulic conductances. Under unlimited light conditions, a within-crown trade-off between A _n/g _s and K _T becomes apparent. Differences in stomatal conductance between the detached shoots from sunlit and shaded canopy layers were largely attributable to the variation in stomatal morphology; significant relationships were established with characteristics combining stomatal size and density (relative stomatal surface, stomatal pore area index). Stomatal morphology is very likely involved in long-term adjustment of photosynthetic WUE.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Stomatal and photosynthetic responses during sun/shade transitions in subalpine plants: influence on water use efficiency

Summary Different response patterns in net photosynthesis (A) leaf conductance (g) and water use efficiency (WUE= a/transpiration) in three subalpine plants occurred during experimental sun/shade transitions that simulated natural cloudcover. In Frasera speciosa Dougl., a large-leaved herb characteristic of open sites, g was relatively insensitive to transitions in irradiance and variations in A. However, large decreases in leaf temperature resulted in reduced transpiration during shade intervals and relatively constant WUE throughout the experimental sun/shade regime. In the understory herb, Arnica cordifolia Hook., patterns of A and g were similar during sun/shade transitions, but WUE was substantially reduced compared to steady-state levels. A third, somewhat intermediate pattern of A, g, and WUE was found in Artemisia tridentata L., an open site shrub. Higher intercellular CO₂ values in A. tridentata suggested that internal, cellular limitations to A were high relative to stomatal limitations in this shrub when compared to the herbaceous species.     

Differential response of leaf gas exchange to enhanced ultraviolet-B (UV-B) radiation in three species of herbaceous climbingplants

We measured diurnal changes in photosynthetic rate, transpiration rate, stomatal conductance and water use efficiency in three species of herbaceous climbing plants (Luffa cylindrica, Trichosanthes kirilowii and Dioscorea opposita) exposed to two intensities of UV-B radiation: 3.0 μw cm^?2 (R1) and 8.0 μw cm^?2 UV-B (R2) radiation under ambient growth conditions. Responses differed per species and per treatment. In Luffa all values increased compared to the Control in both treatments, except for stomatal conductance in R2. In Trichosanthes photosynthetic rates and water use efficiency increased, while the transpiration rates decreased under both treatments, and stomatal conductance was lower in R1. In Dioscorea photosynthetic rates and water use efficiency decreased under both treatments, while the transpiration rates and stomatal conductance increased. The results suggested that to some extent increased UV-B radiation was beneficial to the growth of L. cylindrica and T. kirilowii, but detrimental to D. opposita.     

Flooding and drought tolerance in seeds and seedlings of two Mora species segregated along a soil hydrological gradient in the tropical rain forest of Guyana

Mora excelsa and M. gonggrijpii are well segregated along a soil hydrological gradient. M. excelsa is positively associated with soil hydromorphic characteristics such as gley, mottling and groundwater within 1.20 m, whereas M. gonggrijpii is negatively associated with these characteristics. Growth and mortality of artificially installed seedlings were studied in both species in occasionally flooded forest and dryer uphill forest. In a moderate year (1992, no pronounced flooding, no drought), there was no difference between the two species in growth or mortality in the two forest types. M. gonggrijpii was larger in both forest types. Flooding tolerance of seeds and seedlings were studied under controlled water regimes. Seeds of M. gonggrijpii appeared to be very intolerant to flooding, since germination in this species dropped to 50% after only 11 days of flooding. Seeds of M. excelsa floated and 80% of the seeds were viable after as much as 50 days of flooding. Artificially submerged seeds of the latter species had an intermediate survival response. Flooding in seedlings resulted in cessation of growth in both species. Mortality was nil in most treatments, but all M. gonggrijpii individuals died after a treatment of 8 weeks of continuous flooding. Drought tolerance of seedlings of M. excelsa and M. gonggrijpii was studied in a drying-out experiment. Seedlings of both species of approximately equal size differed widely in a number of characteristics: total leaf area, leaf dry weight, leaf thickness, leaf size and leaf area ratio (LAR) were all larger in M. gonggrijpii, while stomatal density and specific leaf area (SLA) were smaller in this species. Seedlings did not differ in stem hydraulic conductivity. M. excelsa showed lower osmotic potential at full hydration. Turgor potential loss points were not nearly approached in the forest during the middle of the dry season in either species. M. gonggrijpii had much lower stomatal conductance than M. excelsa, due to lower stomatal density. Boundary layer conductance was of the same magnitude as stomatal conductance, especially in the morning. In a drying-out experiment, total plant transpiration was higher in M. gonggrijpii, as the lower conductance observed in this species was compensated for by its larger leaf area. M. gonggrijpii was able to extract water from dryer soils than M. excelsa and may be able to utilize its higher leaf water content under moderate drought in the forest understorey.     

Light as a regulator of structural and chemical leaf defenses against insects in two Prunus species

Light is a key factor influencing competition between species, and the mechanisms by which trees overcome insect outbreaks can be associated with alternation of the leaves structure, which then prevent or promotes their susceptibility to herbivores. It was predicted that leaf tissue anatomy would likely be different in sun and shade leaves, with a gradual decline of leaves resistance coupled with reduction of accessible light. We quantified anatomical patterns and the distribution of defence compounds (phenols, total tannins, catechol tannins) within heavily grazed leaves of Prunus padus, native in Europe and Prunus serotina, an invasive to Central Europe. Both species were strongly attacked by folivorous insects when shrubs grew in the shade. In the sun, however only P. padus leaves were grazed, but P. serotina leaves were almost unaffected. We identified that anatomical characteristics are not linked to different P. padus and P. serotina leaf vulnerability to insects. Furthermore, the staining of defence compounds of P. serotina leaves grown in full sun revealed that the palisade mesophyll cells had a higher content of phenolic compounds and catechol tannins. Thus, our results indicate that a specific distribution of defence compounds, but not the anatomical relationships between palisade and spongy mesophyll, may be beneficial for P. serotina growth outside its natural range. The identified pattern of defence compounds distribution is linked to a lower susceptibility of P. serotina leaves to herbivores, and is associated with its invasiveness. This likely reflects that P. serotina is a stronger competitor than P. padus, especially at high sunlit sites i.e. gaps in the forest.     

RAPID LEAF MOVEMENT,MICROCLIMATE, AND WATER RELATIONS OF TWO TEMPERATE LEGUMES IN THREE CONTRASTING HABITATS

Diurnal heliotropic leaf movements, microclimate, stomatal conductance and leaf water potential of two leguminous species, Strophostyles helvola and Amphicarpa bracteata, were measured in three different habitats over two growing seasons. The habitats occurred along an environmental gradient from an open, sandy beach to a closed canopy deciduous forest understory. At the beach site, heliotropism in S. helvola resulted in higher irradiances in morning and afternoon hours and lower irradiances during midday periods compared to an horizontal leaf. In an exposed forest site A. bracteata responded within minutes to penetration of the direct solar beam by orienting its leaves to steep angles. In contrast, plants in closed canopy locations showed little diurnal leaf movement. The combined results of leaf energy budget calculations and plant physiological responses suggest that ecological ramifications of these movements vary with habitat. We hypothesize that heliotropism in open habitats increases water use efficiency and maximizes carbon returns on plant investment in photosynthesis. Conversely, we hypothesize that leaf movements in understory habitats represent a morphological mechanism to avoid thermal damage, photoinhibition of the photosynthetic apparatus, and water stress associated with high irradiances.     

RESPONSE OF ANDROPOGON GERARDII (POACEAE) TO FIRE-INDUCED HIGH VS. LOW IRRADIANCE ENVIRONMENTS IN TALLGRASS PRAIRIE: LEAF STRUCTURE AND PHOTOSYNTHETIC PIGMENTS

Photosynthetic pigments and several structural characteristics were measured in leaves of Andropogon gerardii from tallgrass prairie populations in an unburned, low-irradiance site and a burned, high-irradiance site to determine if these species displayed sun/shade differences similar to those documented in forest species. Early in the growing season, leaves of A. gerardii in the low-irradiance, unburned site had significantly lower stomatal density, pore length, and conductance, as well as specific leaf mass and thickness than leaves from the high-irradiance, burned site. Moreover, the chlorophyll a:b ratio, carotenoid content, and bundle sheath-vascular complex area were significantly lower, but chlorophyll content (mass/mass) was greater in leaves in unburned vs. burned sites. These differences are consistent with sun/shade adaptations reported for forest understory plants and may contribute to the low productivity of A. gerardii in unburned tallgrass prairie.     

Effect of canopy gap light environment on evaporative load and stomatal conductance in the temperate forest understory herb Aster macrophyllus (Asteraceae)

Aster macrophyllus, a temperate forest understory species of the northeastern United States, inhabits a broad range of light habitats. Plants receiving several minutes of direct sun in canopy gap and forest edge habitats occasionally wilt, a response indicative of water stress. We compared two alterative scenarios for patterns of evaporative load and stomatal conductance for plants in large (0.15 ha) tree canopy gaps and small (3 m²) herbaceous subcanopy gaps: 1) evaporative loads are typically moderate and stomatal conductance is largely governed by light intensity; or 2) evaporative loads are often substantial, mandating stomatal closure to prevent excessive transpiration. In all cases evaporative loads were elevated by light intensity above 25% of full sun. This was accompanied by substantial stomatal closure. Transitions from low to moderate light intensity (<13% full sun) caused little increase in leaf evaporative load, and stimulated increases in stomatal conductance. Very brief periods of high light also stimulated stomatal opening. Light environments in the small herbaceous subcanopy gaps differ greatly in their patterns of evaporative load from day to day.     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

Influence of water stress and low irradiance on morphological and physiological characteristics of <Emphasis Type="Italic">Picea asperata</Emphasis> seedlings

The combined effects of water stress (WS) and low irradiance (LI) on growth, photosynthesis, osmotic adjustment, and lipid peroxidation were studied in dragon spruce (Picea asperata Mast.) seedlings grown under two water treatments (well watered, 100 % of field capacity, and water stressed, 30 % of field capacity) and two irradiances (HI, 100 % of full sunlight and LI, 15 % of full sunlight). WS reduced growth, chlorophyll (Chl) a and b contents, net photosynthetic rate, transpiration rate, stomatal conductance, and effective quantum yield of photosystem 2 (Y) but increased free proline and malondialdehyde contents. LI increased Chl contents and decreased Y, photochemical quenching (q_P), and non-photochemical quenching (q_N) under both water treatments. Hence the seedlings in the understory were more sensitive to drought than to LI.     

Stomatal and nonstomatal limitations of photosynthesis in relation to the drought and shade tolerance of tree species in open and understory environments

 Light saturated photosynthesis (A) in field saplings of shade tolerant, intermediate, and intolerant tree species was analyzed for stomatal and nonstomatal limitations to test differences between species and sun and shade phenotypes during drought. Throughout the study, photosynthesis was highest and mesophyll limitations of A (L_m) lowest in the intolerant species in both open and understory habitats. The shade tolerant species exhibited the only drought-related decreased A and increased L_m in the open, and the greatest drought-related decreased A and increased L_m in the understory. Few species exhibited significant habitat or drought-related differences in stomatal conductance to CO₂ (g_c), but even slight decreases in g_c during drought were associated with large increases in stomatal limitations to A (L_g). Combined changes in L_m and L_g resulted in increased relative stomatal limitation to A (l _g) in several species during drought. Nevertheless, the overall lack of stomatal closure allowed for nonstomatal limitations to play a major role in reduced A during drought. Higher leaf N was associated with shallower slope of the l _g versus g_c relationship, an indication of greater A capacity. Photosynthetic capacity tended to be greater in the intolerant species than the tolerant species, and it tended to decrease during drought primarily in the shade tolerant species in the understory. Findings in the literature suggest that carbon reduction reactions may be more susceptible to drought than photosynthetic light reactions. If so, reduced carbon reduction capacity of shade tolerant species or shade phenotypes may predispose them to drought conditions, which suggests a mechanism behind the well-recognized tradeoff between drought tolerance and shade tolerance of temperate tree species. Received: 20 October 1995 / Accepted: 20 February 1996