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Changes in soil properties after establishment of Artemisia halodendron and Caragana microphylla on shifting sand dunes in semiarid Horqin Sandy Land, northern China.

In the semiarid Horqin sandy land of northern China, establishment of artificial sand-fixing shrubs on desertified sandy lands is an effective measure to control desertification and improve the regional environment. Caragana microphylla Lam. and Artemisia halodendron Turcz. ex Bess. are two of the dominant native shrub species, which are adapted well to windy and sandy environments, and thus, are widely used in revegetation programs to control desertification in Horqin region. To assess the effects of artificially planting these two shrub species on restoration of desertified sandy land, soil properties and plant colonization were measured 6 years after planting shrubs on shifting sand dunes. Soil samples were taken from two depths (0-5 cm and 5-20 cm) under the shrub canopy, in the mid-row location (alley) between shrub belts, and from nonvegetated shifting sand dune (as a control). Soil fine fractions, soil water holding capacity, soil organic C and total N have significantly increased, and pH and bulk density have declined at the 0-5-cm topsoil in both C. microphylla and A. halodendron. At the 5-20 cm subsurface soil, changes in soil properties are not significant, with exception of bulk density and organic C concentration under the canopy of A. halodendron and total N concentration under the canopy of C. microphylla. Soil amelioration processes are initiated under the shrub canopies, as higher C and N concentrations were found under the canopies compared with alleys. At the same time, the establishment of shrubs facilitates the colonization and development of herbaceous species. A. halodendron proved to have better effects in fixing the sand surface, improving soil properties, and restoring plant species in comparison to C. microphylla.

Aluminum Silicates↗

Short-term variations in delta(13)C of ecosystem respiration reveals link between assimilation and respiration in a deciduous forest.

We present a comprehensive dataset of hourly, daily, and monthly measurements of carbon isotope measurements of CO(2) in canopy air from a temperate deciduous forest with the aim to identify the relevance of short-term variations in the isotopic signature of ecosystem respiration (delta(13)C(R)) and to understand its underlying physiological processes. We show that during daytime low vertical mixing inside the canopy can lead to decoupling of the air in the lower and upper canopy layer resulting in large spatial variation of delta(13)C in CO(2) of canopy air. Intercept of Keeling Plots also showed large temporal variation (3.8 per thousand) over the course of the day demonstrating that intercepts can differ between day and night and suggesting that choosing the right time for sampling is essential to capture the isotopic signature of ecosystem respiration (delta(13)C(R)). delta(13)C(R) as obtained from night-time measurements showed large variation of up to 2.65 per thousand on a day-to-day basis, which was similar to the observed variation of delta(13)C(R) over the seasonal cycle (3.08 per thousand). This highlights the importance of short-term physiological processes within ecosystems for the isotopic composition of CO(2) in the atmosphere, not reflected by bulk plant and soil organic samples. At daily and monthly time scales, delta(13)C(R) increased with increasing ratio of vapour pressure deficit to photosynthetically active radiation, measured 4-5 days before. This suggests that ecosystem respiration was isotopically linked to assimilation. Furthermore, assimilates recently fixed in the canopy seem to form a labile carbon pool with a short mean residence time that is respired back to the atmosphere after 4-5 days.

Air↗

Spatially variable propagule pressure and herbivory influence invasion of chaparral shrubland by an exotic grass.

Although numerous studies have identified mechanisms that either resist or facilitate biological invasions, few studies have explicitly tested how resisting and facilitating mechanisms interact to drive invasion success. In California, USA, undisturbed Mediterranean-type shrublands have resisted invasion by the perennial tussock grass Cortaderia jubata. In some cases, however, this resistance has been spectacularly breached even in the absence of large-scale disturbance. I tested the hypothesis that these invasions are facilitated by local reductions in the strength of biotic resistance. I evaluated invasive success using C. jubata seed and seedling additions at different microhabitats: the edge of a chaparral stand, under shrub canopy at different distances from the stand edge, and in canopy gaps within the stand. When left exposed to mammalian herbivores, seedling survivorship decreased sharply from nearly 40% on the stand edge to zero just 10 m into the stand. When transplants were protected from herbivory, however, distance from the edge had no significant influence on transplant survivorship. Seedling emergence was also greater on the edge and in canopy gaps than under the canopy, but these differences were not caused by differences in herbivory. The flux of invasive propagules reaching the soil surface was immense and greater along the edge and within gaps than under the stand canopy. Mirroring these patterns, naturally occurring seedling abundance declined dramatically with distance from the stand edge, and seedlings were far more common within stand gaps than would be expected given gap frequency within the stand. Despite strong biotic resistance to invasion within the stand, the cover of C. jubata has increased 20% over the last 9 years. These results suggest that the relative amount of susceptible edge habitat and the supply of invasive propagules can facilitate invasion even in the face of strong local biotic resistance.

Animals↗

Nitrate sequestration by corticolous macrolichens during winter precipitation events.

Nitrogen is an essential nutrient in the biogeochemistry of forested ecosystems. The influence of canopy lichens on the winter biogeochemistry of nitrate in broadleaved deciduous forests is examined and it is hypothesized that nitrate sequestration will not differ between winter precipitation events. Rejection of this hypothesis would mean that meteorological conditions of winter precipitation events have a detectable influence on nitrate sequestration by canopy lichens and nitrate input to the forest floor. Canopy lichens of the genus Parmelia were found to influence winter nitrate stemflow inputs to forest soils differentially. Epiphytic lichens on an individual Carya glabra Mill. (pignut hickory) canopy tree, centrally located within the stand of an open deciduous forest, actively sequestered nitrate leached from the tree's woody frame, lowering aqueous stemflow inputs at the tree base. The quantities of nitrate sequestered by corticolous lichens during the 2 February 1999 mixed-precipitation event were significantly greater than those during all other precipitation events examined. Greater rates of nitrate uptake during the 2 February 1999 event may be attributed to (1) its intermediate rain intensity, which would have soaked the lichen thalli in a nutrient-rich bath, and (2) an air temperature range between -2 degrees C and 8 degrees C that would have increased viscosity and surface tension of stemflow drainage, thereby decreasing stemflow velocity and increasing the contact time of stemflow water on the lichen thalli. Other precipitation events were either too cold to promote metabolic activity by canopy lichens or too warm and intense for an optimal contact time of stemflow with lichen thalli, resulting in lower quantities of nitrate sequestered. Meteorological conditions of winter precipitation events have been documented to influence sequestration of nitrate by corticolous lichens and decrease aqueous stemflow inputs to the forest floor of broadleaved deciduous forests.

Ecosystem↗

Forest gap dynamics and the Ising model.

The vegetation height in forest ecosystems is spatially heterogeneous. Canopy gaps (sites with low vegetation) are formed by treefalls, and they recover to canopy sites (with high vegetation) either by growth of small trees or by branch extension of surrounding trees. The dynamics of canopy gaps have been studied using a spatial Markov chain with nearest neighbor interaction. (1) If the canopy recovery rate is constant and if the gap formation rate for a site increases exponentially with the number of neighboring gap sites, the equilibrium distribution is the same as the one generated by the Ising model in statistical mechanics. Here, we extend the equivalence to the situation in which both the gap formation and canopy recovery depend on the neighborhood, as shown in recent forest data. (2) We develop a statistical test of whether a given spatial pattern is a random sample from the Ising model. The test is based on the conditional probability of configurations on a partial lattice. We apply the method to vegetation height data from the Ogawa forest reserve, Japan, measured on a 5x5 m grid in 1976, 1981, 1986, and 1991. The spatial pattern of the original forest data deviates significantly from the Ising model. We examine whether a larger sampling distance or the removal of the effects of the topography can reduce this deviation.

Ecosystem↗

Modern pollen-representation of some boreal species on islands in a large lake in Canada.

Studies of pollen source areas of closed-canopy sites are contradictory. Some authors found that closed-canopy sites mainly collect local pollen while others found more distant sources. This dichotomy might stem from the use of canopies of varying degrees of closure, and from variations in the pollen productivity of the local vegetation and the background pollen rain. Here, 30 islands were used to evaluate the pollen sources of closed-canopy sites. We compared pollen with the forest inventory in three quadrat sizes: 100, 400m(2) and on the whole island. Regression analyses showed that most pollen of Picea spp., Pinus spp., and Betula spp. comes from within the 400m(2) quadrat. Abies balsamea and Thuja occidentalis showed no relationship with vegetation in any of the quadrats considered, suggesting a more regional source. Insularity and island size are important factors influencing the pollen source area; correlations were stronger on islands located 120050ha. These results suggest that closed-canopy sites on islands may be useful in stand-level vegetation history reconstruction through pollen analysis, but that caution must be exercised in separating the local and regional signals.

Journal Article↗

Aerial contamination of fruit through wet deposition and particulate dry deposition.

The current state of knowledge concerning the processes and pathways which lead to the contamination of fruit crops by non-gaseous airborne contaminants is reviewed. Given the wide range of fruit canopies which occur, it is necessary to look for generic factors which affect the contribution of each of these processes and pathways to distribution through the canopy, losses from the canopy back to the atmosphere, and the fate of particle-bound substances once attached to the canopy. This latter stage represents perhaps the greatest source of uncertainty in determining levels of contamination. For wet deposition, the controlling factors appear to be the ability of the canopy surface to store precipitated water, and the interaction of the contaminant species with the leaf cuticle, which appears to act as an ion exchange medium, selectively accumulating certain ionic species. Where possible, models and parameterisations are provided for these stages. A summary is given of current data requirements for those cases where this is not possible.

Air Movements↗

Drought stress and carbon uptake in an Amazon forest measured with spaceborne imaging spectroscopy.

Amazônia contains vast stores of carbon in high-diversity ecosystems, yet this region undergoes major changes in precipitation affecting land use, carbon dynamics, and climate. The extent and structural complexity of Amazon forests impedes ground studies of ecosystem functions such as net primary production (NPP), water cycling, and carbon sequestration. Traditional modeling and remote-sensing approaches are not well suited to tropical forest studies, because (i) biophysical mechanisms determining drought effects on canopy water and carbon dynamics are poorly known, and (ii) remote-sensing metrics of canopy greenness may be insensitive to small changes in leaf area accompanying drought. New spaceborne imaging spectroscopy may detect drought stress in tropical forests, helping to monitor forest physiology and constrain carbon models. We combined a forest drought experiment in Amazônia with spaceborne imaging spectrometer measurements of this area. With field data on rainfall, soil water, and leaf and canopy responses, we tested whether spaceborne hyperspectral observations quantify differences in canopy water and NPP resulting from drought stress. We found that hyperspectral metrics of canopy water content and light-use efficiency are highly sensitive to drought. Using these observations, forest NPP was estimated with greater sensitivity to drought conditions than with traditional combinations of modeling, remote-sensing, and field measurements. Spaceborne imaging spectroscopy will increase the accuracy of ecological studies in humid tropical forests.

Brazil↗

Modelling plant responses to elevated CO2: how important is leaf area index?

BACKGROUND AND AIMS: The problem of increasing CO(2) concentration [CO(2)] and associated climate change has generated much interest in modelling effects of [CO(2)] on plants. While variation in growth and productivity is closely related to the amount of intercepted radiation, largely determined by leaf area index (LAI), effects of elevated [CO(2)] on growth are primarily via stimulation of leaf photosynthesis. Variability in LAI depends on climatic and growing conditions including [CO(2)] concentration and can be high, as is known for agricultural crops which are specifically emphasized in this report. However, modelling photosynthesis has received much attention and photosynthesis is often represented inadequately detailed in plant productivity models. Less emphasis has been placed on the modelling of leaf area dynamics, and relationships between plant growth, elevated [CO(2)] and LAI are not well understood. This Botanical Briefing aims at clarifying the relative importance of LAI for canopy assimilation and growth in biomass under conditions of rising [CO(2)] and discusses related implications for process-based modelling. MODEL: A simulation exercise performed for a wheat crop demonstrates recent experimental findings about canopy assimilation as affected by LAI and elevation of [CO(2)]. While canopy assimilation largely increases with LAI below canopy light saturation, effects on canopy assimilation of [CO(2)] elevation are less pronounced and tend to decline as LAI increases. Results from selected model-testing studies indicate that simulation of LAI is often critical and forms an important source of uncertainty in plant productivity models, particularly under conditions of limited resource supply. CONCLUSIONS: Progress in estimating plant growth and productivity under rising [CO(2)] is unlikely to be achieved without improving the modelling of LAI. This will depend on better understanding of the processes of substrate allocation, leaf area development and senescence, and the role of LAI in controlling plant adaptation to environmental changes.

Biomass↗

Use of infrared thermography for monitoring stomatal closure in the field: application to grapevine.

This paper reviews and discusses strategies for the use of thermal imaging for studies of stomatal conductance in the field and compares techniques for image collection and analysis. Measurements were taken under a range of environmental conditions and on sunlit and shaded canopies to illustrate the variability of temperatures and derived stress indices. A simple procedure is presented for correcting for calibration drift within the images from the low-cost thermal imager used (SnapShot 225, Infrared Solutions, Inc.). The use of wet and dry reference surfaces as thresholds to eliminate the inclusion of non-leaf material in the analysis of canopy temperature is discussed. An index that is proportional to stomatal conductance was compared with stomatal measurements with a porometer. The advantages and disadvantages of a possible new approach to the use of thermal imagery for the detection of stomatal closure in grapevine canopies, based on an analysis of the temperature of shaded leaves, rather than sunlit leaves, are discussed. Evidence is presented that the temperature of reference surfaces exposed within the canopy can be affected by the canopy water status.

Algorithms↗

Seasonal development of leaf area in a young, widely spaced Pinus radiata D. Don stand.

Measurements of needle elongation and needle death were made at two-week intervals during a year on 250 branch units spread throughout the crowns of six trees (three high-pruned, three low-pruned) in a widely spaced, 6- to 7-year-old Pinus radiata D. Don plantation in New Zealand. The trees were well supplied with nutrients and water. During the year, mean tree height increased by 1.2 m and the cross-sectional area of stem below the green crown (used to predict leaf area) for the average tree increased from 8.7 x 10(3) to 13.9 x 10(3) mm(2). The increase in stem cross-sectional area occurred throughout the year except for two months in early winter (May and June). Elongation of Age 0 needles began in Spring (October), continued through summer, and the mean date for 95% completion of elongation was in autumn (early May), approximately 200 days after elongation began. Mean maximum needle length in the canopy decreased with increasing branch order and was 136 and 94 mm for Order 1 and Order 3 units, respectively. Needle elongation was related to thermal time, using growing degree days with a base temperature of 6 degrees C. The mean maximum rate of needle elongation in the canopy was 0.11 m ( degrees C day)(-1) and this occurred in early summer (mid-December), 47 days after elogation started. Maximum needle length and the rate of elongation increased, and the time taken to reach 95% elongation decreased with increasing height in the canopy. A smaller autumn flush of needles started in summer (January) and the needles elongated linearly at a mean rate of 0.07 mm ( degrees C day)(-1) until the end of the growing season when temperatures fell below the base value. At the end of the year, the mean length of needles from the autumn flush was 66 mm. The density of needles did not change with height in the canopy and there were no significant changes seasonally. The mean density values for Age 1 and Age 0 needles were 336 and 286 kg m(-3), respectively. Dry weight per unit length did not change seasonally for Age 1 needles, but the mean values for Order 1 and Order 2 needles were 0.32 and 0.23 mg mm(-1), respectively. Dry weight per unit length for Age 0 needles increased during the growing season (October to February). This was particularly apparent for Order 1 needles where the mean value increased from 0.12 to 0.25 mg mm(-1). Death of Age 1 needles started in midsummer (mean date was January 24) which coincided with the time of maximum elongation of Age 0 needles, but there were differences in timing among individual trees. The mean rate of death was 1.7% day(-1) and the mean duration was 59 days. Leaf area index for the stand increased from 3.2 to 5.3 (all-surfaces basis) during the year. At the end, 92% of the leaf area had grown during the year and 60% of it was on Order 2 branch units. The models for needle elongation and needle death were used to scale up to seasonal changes in canopy leaf area index. Leaf area index peaked with increasing Age 0 leaf area to 6.1 in summer (January), fell to 4.7 in March as Age 1 needles died, then increased slightly again. The seasonal dynamics are consistent with the hypothesis of Cannell (1989) that the timing of maximum leaf area index within a year is optimal for maximizing biomass production.

Journal Article↗

Leaf water potential and stomatal conductance in Quercus pyrenaica Willd. forests: vertical gradients and response to environmental factors.

Two permanent sampling sites were selected at the two extremes of a rainfall gradient in natural forests of Quercus pyrenaica Willd. located in the Sierra de Gata (Salamanca Province, Spain). During 1991 and 1992, diurnal courses of transpiration rate (E), stomatal conductance (g(s)) and leaf water potential (Psi(l)) were studied at different levels in the tree canopy. Leaf water potential fell from a high value in the early morning to a minimum at midday and then increased again in the afternoon. Values of Psi(l) in the early morning and at midday were lower at Fuenteguinaldo (dry site) than at Navasfrías (wet site) and were related to soil water availability. Stomatal conductance increased during the morning and maximum g(s) occurred about 3 h before Psi(l) was at its lowest value. The highest values of g(s) were found at Navasfrías during July. The lowest Psi(l) values were found in the upper parts of the canopy and differences among canopy levels were greatest when soil water availability was low. In August, minimum values of Psi(l) differed among canopy levels by 1.0 MPa at Navasfrías and by 0.8 MPa at Fuenteguinaldo. The maximum difference in g(s) among canopy levels was about 150 mmol m(-2) s(-1) at Navasfrías and about 300 mmol m(-2) s(-1) at Fuenteguinaldo. The results indicate a nonconservative pattern of water use in Quercus pyrenaica.

Journal Article↗

Field measurements of isoprene emission from trees in response to temperature and light.

The atmospheric hydrocarbon budget is important for predicting ozone episodes and the effects of pollution mitigation strategies. Isoprene emission from plants is an important part of the atmospheric hydrocarbon budget. We measured isoprene emission capacity at the bottom, middle, and top of the canopies of a white oak (Quercus alba L.) tree and a red oak (Quercus rubra L.) tree growing adjacent to a tower in the Duke University Forest. Leaves at the top of the white oak tree canopy had a three- to fivefold greater capacity for emitting isoprene than leaves at the bottom of the tree canopy. Isoprene emission rate increased with increasing temperature up to about 42 degrees C. We conclude that leaves at the top of the white oak tree canopy had higher isoprene emission rates because they were exposed to more sunlight, reduced water availability, and higher temperature than leaves at the bottom of the canopy. Between 35 and 40 degrees C, white oak photosynthesis and stomatal conductance declined, whereas red oak (Quercus rubra) photosynthesis and stomatal conductance increased over this range. Red oak had lower rates of isoprene emission than white oak, perhaps reflecting the higher stomatal conductance that would keep leaves cool. The concentration of isoprene inside the leaf was estimated with a simplified form of the equation used to estimate CO(2) inside leaves.

Journal Article↗

Rainfall interception and drop size-development and calibration of the two-layer stochastic interception model.

This paper reviews the development of the stochastic interception model from the original, single-layer, drop-size-dependent model to the two-layer model that recognizes that vegetation canopies are wetted through both the primary impact of raindrops to the top layer of the canopy and secondary impacts from drops falling from the vegetation to lower layers of the canopy. It is shown that drop volumes of primary raindrops can be calculated from the Marshall-Palmer distribution and drop volumes of secondary drops can be estimated from disdrometer measurements of the characteristic volume appropriate to the particular vegetation species. It is recognized that, in addition to the volume-dependent stochastic wetting effect, there is also another drop-size-dependent wetting effect that is related to the kinetic energy of the raindrops, which reduces the maximum storage that can be achieved on the canopy. The predicted wetting functions for canopies of different density are described and compared with observations made with the use of a rainfall simulator. It is also shown that the species-dependent model parameters can be determined from measurements made with the rainfall simulator. The improved performance of the model compared with conventional interception models is demonstrated for a tropical forest in Sri Lanka. Application of the two-layer model may explain why interception losses from coniferous, fine-leaved forests in the temperate, low-intensity rainfall climate of the uplands of the U.K. are among the highest in the world, whereas interception losses from tropical broad leaved forest in high-intensity rainfall climates of the tropics are among the lowest.

Journal Article↗

Foliar temperature-respiration response functions for broad-leaved tree species in the southern Appalachians.

We measured leaf respiration in 18 eastern deciduous forest tree species to determine if there were differences in temperature-respiration response functions among species or among canopy positions. Leaf respiration rates were measured in situ and on detached branches for Acer pensylvanicum L., A. rubrum L., Betula spp. (B. alleghaniensis Britt. and B. lenta L.), Carya glabra (Mill.) Sweet, Cornus florida L., Fraxinus spp. (primarily F. americana L.), Liriodendron tulipifera L., Magnolia fraseri Walt., Nyssa sylvatica Marsh., Oxydendrum arboreum L., Platanus occidentalis L., Quercus alba L., Q. coccinea Muenchh., Q. prinus L., Q. rubra L., Rhododendron maximum L., Robinia psuedoacacia L., and Tilia americana L. in the southern Appalachian Mountains, USA. Dark respiration was measured on fully expanded leaves at 10, 15, 20, 25, and 30 degrees C with an infrared gas analyzer equipped with a temperature-controlled cuvette. Temperature-respiration response functions were fit for each leaf. There were significant differences in response functions among species and by canopy position within species. These differences were observed when respiration was expressed on a mass, nitrogen, or area basis. Cumulative nighttime leaf respiration was calculated and averaged over ten randomly selected nights for each leaf. Differences in mean cumulative nighttime respiration were statistically significant among canopy positions and species. We conclude that effects of canopy position and species on temperature-respiration response functions may need to be considered when making estimates of whole-tree or canopy respiration.

Journal Article↗

Modeling the influence of temperature on monthly gross primary productivity of sugar maple stands.

A bottom-up and a top-down model were used to estimate the effect of temperature on monthly gross primary productivity (GPP) of sugar maple (Acer saccharum Marsh.). The bottom-up model computed canopy photosynthesis at an hourly time step from detailed physiological sub-models of leaf photosynthesis and stomatal conductance. Leaf mass per area was used as a covariable to integrate photosynthesis through the canopy. The top-down model used a radiation-use efficiency coefficient to relate canopy gross photosynthesis to absorbed photosynthetically active radiation at a monthly time step. The parameters of the top-down model were estimated from simulations with the bottom-up model. Forty single-year simulations were made using records of daily maximum and minimum temperatures from weather stations selected within the natural range of sugar maple in the province of Québec, Canada. Leaf area index was randomly varied between 4 and 10. Within a broad range of values, temperature had a minor effect on predicted monthly canopy-level GPP and its contribution to explaining the variability of GPP was low, both through its direct effect on photosynthetic processes (1.1%), and indirectly through the effect of relative humidity on stomatal conductance (4.0%). This result was unchanged when key parameters relating photosynthesis to temperature and stomatal conductance to atmospheric humidity were changed in the bottom-up model. An increase in time step from hourly to monthly resulted in a downward shift in the optimum temperature range for photosynthesis, from 30 degrees C for a leaf at saturating irradiance to 22 degrees C for the canopy at a monthly time scale.

Journal Article↗

Structural differences and functional similarities between two sugar maple (Acer saccharum) stands.

The spatially inexplicit or functional multilayer models used to predict canopy transpiration or photosynthesis are based on the assumption that closed stands show less functional variability than structural variability, because foliage tends to arrange itself in space to optimize the capture of light. To validate this assumption, we compared the structural and functional properties, and the measured and modeled transpiration fluxes of two sugar maple (Acer saccharum Marsh.) stands of comparable leaf mass but differing in height and diameter distributions. One stand was characterized by a well-developed single-layer canopy, whereas the other stand had a multilayered canopy and a stem diameter distribution of the classical inverse-J shape. Stand differences in height and diameter distribution, and canopy gap fraction, were highly significant. There were minor but significant differences in leaf mass and leaf mass per unit leaf area (LMA) distributions. We found no differences in tree-level relationships between basal area and either transpiration flux or sapwood area. We compared measurements of stand transpiration with transpiration estimates obtained from a multilayer gas exchange model, in which only the nonspatial inputs, leaf area index and LMA frequency distribution described stand structure. For both stands, modeled values of daily transpiration closely followed measured values (r(2) = 0.94). These results support use of the nonspatially explicit approach to estimating canopy gas exchange, especially if the intent is to scale-up to larger portions of the landscape.

Acer↗

Stomatal and non-stomatal limitations to photosynthesis in four tree species in a temperate rainforest dominated by Dacrydium cupressinum in New Zealand.

We assessed the relative limitations to photosynthesis imposed by stomatal and non-stomatal processes in Dacrydium cupressinum Lamb. (Podocarpaceae), which is the dominant species in a native, mixed conifer-broad-leaved rainforest in New Zealand. For comparison, we included three co-occurring broad-leaved tree species (Meterosideros umbellata Cav. (Myrtaceae), Weinmannia racemosa L.f. (Cunoniaceae) and Quintinia acutifolia Kirk (Escalloniaceae)) that differ in phylogeny and in leaf morphology from D. cupressinum. We found that low foliage phosphorus content on an area basis (P(a)) limited light-saturated photosynthesis on an area basis (A(sat)) in Q. acutifolia. Depth in the canopy did not generally affect A(sat) or the relative limitations to A(sat) because of stomatal and non-stomatal constraints, despite reductions in the ratio of foliage mass to area, foliar nitrogen on an area basis (N(a)) and P(a) with depth in the canopy. In the canopy-dominant conifer D. cupressinum, A(sat) was low, consistent with low values of the maximum rate of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) carboxylation (V(cmax)). In comparison, the A(sat) response of the three broad-leaved tree species was quite variable. Although A(sat) was high in the canopy-dominant M. umbellata, it was low in the sub-canopy trees W. racemosa and Q. acutifolia. Relative stomatal limitation to photosynthesis was more pronounced in W. racemosa (40%) than in the other three species (28-33%). Despite differences in degree, non-stomatal limitation to A(sat) predominated in all tree species.

Chlorophyll↗