Flows through forest canopies in complex terrain

Recent progress on boundary layer flow within and above tall forest canopies in complex terrain is reviewed from the perspective of developing methods to interpret carbon dioxide fluxes from tower measurements in real terrain. Two examples of complex terrain are considered in detail: a forest edge, which exemplifies nonuniform forests, and hilly terrain, which can lead to drainage currents at night. Dynamical arguments show that, when boundary layer winds approach a forest edge, the mean wind adjusts on a length scale of approximately 3L(c), where L(c) is the canopy drag length scale, which depends inversely on the leaf area density of the forest. Over a further distance that also scales on L(c), turbulence in the flow adjusts, and the mixing and transport in the canopy approaches the homogeneous limit. Even low hills change the neutral flow within and above the forest canopy substantially. When the canopy is tall, pressure gradients drive flow up both the upwind and downwind slopes of the hill, leading to an ejection of air out of the top of the canopy just downwind of the crest. This flow at the crest can then advect scalar out of the top of the forest, leading to large variations in the flux of scalar across the hill. At night, when the air near the ground cools and becomes stably stratified, turbulence within the canopy can collapse, even when the flow above the canopy remains turbulent. This leads to a decoupling of the air motions within the canopy from those above. The air above the canopy can then continue to pass up and over the hill, as it does in the neutral case, but at the same time, air within the canopy drains down the hill slopes as drainage currents. These analyses will help us understand when flux towers are reliably measuring the net ecosystem exchange and suggest ways of correcting the flux tower data in more complex situations.     

The structure of turbulence near a tall forest edge: the backward-facing step flow analogy revisited

Flow disturbances near tall forest edges are receiving significant attention in diverse disciplines including ecology, forest management, meteorology, and fluid mechanics. Current theories suggest that near a forest edge, when the flow originates from a forest into a large clearing, the flow retains its forest canopy turbulence structure at the exit point. Here, we propose that this framework is not sufficiently general for dense forested edges and suggest that the flow shares several attributes with backward-facing step (BFS) flow. Similar analogies, such as rotor-like circulations, have been proposed by a number of investigators, though the consequences of such circulations on the primary terms in the mean momentum balance at the forest clearing edge have rarely been studied in the field. Using an array of three triaxial sonic anemometers positioned to measure horizontal and vertical gradients of the velocity statistics near a forest edge, we show that the flow structure is more consistent with an intermittent recirculation pattern, rather than a continuous rotor, whose genesis resembles the BFS flow. We also show that the lateral velocity variance, v'2, is the moment that adjusts most slowly with downwind distance as the flow exits from the forest into the clearing. Surprisingly, the longitudinal and vertical velocity variances (u'2 and w'2) at the forest edge were comparable in magnitude to their respective values at the center of a large grass-covered forest clearing, suggesting rapid adjustment at the edge. Discussions on how the forest edge modifies the spectra and co-spectra of momentum fluxes, effective mixing length, and static pressure are also presented.     

The role of coherent turbulent structures in explaining scalar dissimilarity within the canopy sublayer

Scalar similarity is widely assumed in models and interpretation of micro-meteorological measurements. However, in the air space within and just above the canopy (the so-called canopy sublayer, CSL) scalar similarity is generally violated. The scalar dissimilarity has been mainly attributed to differences in the distribution of sources and sinks throughout the canopy. Since large-scale coherent structures in the CSL (e.g. double roller and sweep/ejection) arise from the instabilities generated by the interaction between the mean flow and the canopy, they may encode key dynamical features about the production term responsible for the source–sink dissimilarity of scalars. Therefore, it is reasonable to assume that the geometric attributes of coherent structures are tightly coupled to the onset and the vertical extent of scalar dissimilarity within the CSL. Large-eddy simulation (LES) runs were used to investigate the role of coherent structures in explaining scalar dissimilarity among three scalars (potential air temperature, water vapour and $\text{ CO }_2$ concentration) within the CSL under near-neutral conditions for horizontally uniform but vertically varying vegetation leaf area density. It was shown that coherent structures, when identified from the first mode of a novel proper orthogonal decomposition (POD) approach, were able to capture some features of the scalar dissimilarity in the original LES field. This skill was quantified by calculating scalar–scalar correlation coefficients and turbulent Schmidt numbers of the original field and the coherent structures, respectively. However, coherent structures tend to magnify the magnitude of scalar–scalar correlation, particularly in cases where this correlation is already strong. The ability of coherent structures to describe more complex features such as the scalar sweep-ejection cycle was also explored. It was shown that the first mode of the POD does not capture the relative importance of sweeps to ejections in the original LES field. However, the superposition of few secondary coherent structures, derived from higher order POD modes, largely diminish the discrepancies between the original field and the POD expansion.     

Advection and resulting CO2 exchange uncertainty in a tall forest in central Germany

Potential losses by advection were estimated at Hainich Forest, Thuringia, Germany, where the tower is located at a gentle slope. Three approaches were used: (1) comparing nighttime eddy covariance fluxes to an independent value of total ecosystem respiration by bottom-up modeling of the underlying processes, (2) direct measurements of a horizontal CO2 gradient and horizontal wind speed at 2 m height in order to calculate horizontal advection, and (3) direct measurements of a vertical CO2 gradient and a three-dimensional wind profile in order to calculate vertical advection. In the first approach, nighttime eddy covariance measurements were compared to independent values of total ecosystem respiration by means of bottom-up modeling of the underlying biological processes. Turbulent fluxes and storage term were normalized to the fluxes calculated by the bottom-up model. Below a u(*) threshold of 0.6 m/s the normalized turbulent fluxes decreased with decreasing u(*), but the flux to the storage increased only up to values less than 20% of the modeled flux at low turbulence. Horizontal advection was measured by a horizontal CO2 gradient over a distance of 130 m combined with horizontal wind speed measurements. Horizontal advection occurred at most of the evenings independently of friction velocity above the canopy. Nevertheless, horizontal advection was higher when u(*) was low. The peaks of horizontal advection correlated with changes in temperature. A full mass balance including turbulent fluxes, storage, and horizontal and vertical advection resulted in an increase of spikes and scatter but seemed to generally improve the results from the flux measurements. The comparison of flux data with independent bottom-up modeling results as well as the direct measurements resulted in strong indications that katabatic flows along the hill slope during evening and night reduces the measured apparent ecosystem respiration rate. In addition, anabatic flows may occur during the morning. We conclude that direct measurements of horizontal and vertical advection are highly necessary at sites located even on gentle hill slopes.     

Ozon in Waldökosystemen aus atmosphärenchemischer und pflanzenphysiologischer Sicht

The trend of rising ozone concentrations in forest ecosystems and the phytotoxicity of ozone demand a realistic risk assessment according to an internationally accepted and flux-based quality standard. Ozone fluxes within the canopy are influenced by chemical gas-phase reactions with nitrogen oxide and biogenic hydrocarbons and by surface deposition processes. Therefore, a differentiation of the ozone flux within the canopy is needed between stomatal uptake and other transport pathways. The Eddy Covariance technique is the method of choice for the determination of trace gas fluxes relevant for ozone chemistry. This method is also used for stomatal conductance measurements based on evapotranspiration fluxes and for emission measurements of biogenic hydrocarbons by PTR-MS. Although considerably research efforts were directed to canopy measurements in recent years, the underlying processes are not fully understood yet. Thus, major differences occur in the ratios of stomatal ozone uptake, non-stomatal deposition and gas-phase chemistry between different studies. Furthermore, the vertical concentration gradients within the canopy measured at several forest sites are rather inconsistent and the existing deposition models do rarely account for chemical transformation and detoxification processes. Only a simultaneous measurement of all photochemically relevant trace gases, plant physiological parameters at different sites and forest species over entire vegetation periods, and model parameterization according to the measurement results from the experimental sites will contribute to the clarification of the canopy processes and will ensure realistic risk assessments.     

Forest Edges as Nutrient and Pollutant Concentrators: Potential Synergisms between Fragmentation, Forest Canopies, and the Atmosphere

Abstract: Forest fragmentation leads to a dramatic increase in forest edge, and these edges may function as traps and concentrators for wind-borne nutrients and pollutants. We assessed the influence of forest edges on atmospheric deposition and subsequent inputs to the forest floor in deciduous-forest fragments in the eastern United States. To quantify these inputs, we collected throughfall—water that has passed through the forest canopy—from edge and interior zones of forests adjacent to open fields. During the 1995 growing season, atmospheric input (wet and dry deposition) of sulfur to forest edge zones was elevated compared with input to forest interiors. Throughfall fluxes of dissolved inorganic nitrogen and calcium were also greater at edges than interiors. The mean edge increases ranged from 17% to 56% for the nutrients and pollutants we measured. When we manipulated the structure of forest edges by removing all vegetation below half the canopy height, throughfall flux in the edge zone declined sharply and was less than that of the respective interior zone. Changing the vegetation structure of the edge also shifted the zone of highest throughfall flux farther into the interior of the forest. Our data suggest that forest edges can function both as significant traps for airborne nutrients and pollutants from adjoining agricultural or urban landscapes and effective concentrators of below-canopy chemical fluxes. These enhanced fluxes may have cascading effects on soil-nutrient cycling, microbial activity, seedling dominance, and other ecological processes near forest edges.     

The contribution of advective fluxes to net ecosystem exchange in a high-elevation, subalpine forest

The eddy covariance technique, which is used in the determination of net ecosystem CO2 exchange (NEE), is subject to significant errors when advection that carries CO2 in the mean flow is ignored. We measured horizontal and vertical advective CO2 fluxes at the Niwot Ridge AmeriFlux site (Colorado, USA) using a measurement approach consisting of multiple towers. We observed relatively high rates of both horizontal (F(hadv)) and vertical (F(vadv)) advective fluxes at low surface friction velocities (u(*)) which were associated with downslope katabatic flows. We observed that F(hadv) was confined to a relatively thin layer (0-6 m thick) of subcanopy air that flowed beneath the eddy covariance sensors principally at night, carrying with it respired CO2 from the soil and lower parts of the canopy. The observed F(vadv) came from above the canopy and was presumably due to the convergence of drainage flows at the tower site. The magnitudes of both F(hadv) and F(vadv) were similar, of opposite sign, and increased with decreasing u(*), meaning that they most affected estimates of the total CO2 flux on calm nights with low wind speeds. The mathematical sign, temporal variation and dependence on u(*) of both F(hadv) and F(vadv) were determined by the unique terrain of the Niwot Ridge site. Therefore, the patterns we observed may not be broadly applicable to other sites. We evaluated the influence of advection on the cumulative annual and monthly estimates of the total CO2 flux (F(c)), which is often used as an estimate of NEE, over six years using the dependence of F(hadv) and F(vadv) on u(*). When the sum of F(hadv) and F(vadv) was used to correct monthly F(c), we observed values that were different from the monthly F(c) calculated using the traditional u(*)-filter correction by--16 to 20 g C x m(-2) x mo(-1); the mean percentage difference in monthly Fc for these two methods over the six-year period was 10%. When the sum of F(hadv) and F(vadv) was used to correct annual Fc, we observed a 65% difference compared to the traditional u(*)-filter approach. Thus, the errors to the local CO2 budget, when F(hadv) and F(vadv) are ignored, can become large when compounded in cumulative fashion over long time intervals. We conclude that the "micrometeorological" (using observations of F(hadv) and F(vadv)) and "biological" (using the u(*) filter and temperature vs. F(c) relationship) corrections differ on the basis of fundamental mechanistic grounds. The micrometeorological correction is based on aerodynamic mechanisms and shows no correlation to drivers of biological activity. Conversely, the biological correction is based on climatic responses of organisms and has no physical connection to aerodynamic processes. In those cases where they impose corrections of similar magnitude on the cumulative F(c) sum, the result is due to a serendipitous similarity in scale but has no clear mechanistic explanation.     

Large eddy simulation of flow and scalar transport in a vegetated channel

Predicting flow and mass transport in vegetated regions has a broad range of applications in ecology and engineering practice. This paper presents large eddy simulation (LES) of turbulent flow and scalar transport within a fully developed open-channel with submerged vegetation. To properly represent the scalar transport, an additional diffusivity was introduced within the canopy to account for the contribution of stem wakes, which were not resolved by the LES, to turbulent diffusion. The LES produced good agreement with the velocity and concentration fields measured in a flume experiment. The simulation revealed a secondary flow distributed symmetrically about the channel centerline, which differed significantly from the circulation in a bare channel. The secondary circulation accelerated the vertical spread of the plume both within and above the canopy layer. Quadrant analysis was used to identify the form and shape of canopy-scale turbulent structures within and above the vegetation canopy. Within the canopy, sweep events contributed more to momentum transfer than ejection events, whereas the opposite occurred above the canopy. The coherent structures were similar to those observed in terrestrial canopies, but smaller in scale due to the constraint of the water surface.     

Large-eddy simulation of low-level jet-like flow in a canopy

The computational method of Large-Eddy Simulations has been used to study the weak, neutrally stable drainage flow within tree canopies. The computational results show that a secondary velocity maximum that resembles a jet is formed within the canopy under the nocturnal flow conditions. This jet-like flow is important in the analysis and measurements of the net ecosystem-atmosphere exchange (NEE) for carbon dioxide (CO₂). A uniformly distributed, plane source was placed within the canopy in order to simulate the nocturnal production of CO₂. The NEE is calculated as the sum of the integration of the rate of change of the concentration of CO₂ over the computational domain, the vertical turbulent flux measured directly by eddy-covariance (EC) method, and the advection terms, which are not taken into account in the EC method. Numerical results of the velocity and concentration fields, within and above the canopy, are presented and their impact on the CO₂ transport is investigated in detail. The computational results show that 15–20% of NEE is drained out by the advection process under the canopy. The results also show that the turbulent fluctuations in the lateral direction are also significant and may result in 2–5% CO₂ transport.     

Instantaneous transport of a passive scalar in a turbulent separated flow

The results of large-eddy simulations of flow and transient solute transport over a backward facing step and through a 180° bend are presented. The simulations are validated successfully in terms of hydrodynamics and tracer transport with experimental velocity data and measured residence time distribution curves confirming the accuracy of the method. The hydrodynamics are characterised by flow separation and subsequent recirculation in vertical and horizontal directions and the solute dispersion process is a direct response to the significant unsteadiness and turbulence in the flow. The turbulence in the system is analysed and quantified in terms of power density spectra and covariance of velocity fluctuations. The injection of an instantaneous passive tracer and its dispersion through the system is simulated. Large-eddy simulations enable the resolution of the instantaneous flow field and it is demonstrated that the instabilities of intermittent large-scale structures play a distinguished role in the solute transport. The advection and diffusion of the scalar is governed by the severe unsteadiness of the flow and this is visualised and quantified. The analysis of the scalar mass transport budget quantifies the mechanisms controlling the turbulent mixing and reveals that the mass flux is dominated by advection.     

Canopy-wake dynamics and wind sheltering effects on Earth surface fluxes

The atmospheric boundary layer adjustment at the abrupt transition from a canopy (forest) to a flat surface (land or water) is investigated in a wind tunnel experiment. Detailed measurements examining the effect of canopy turbulence on flow separation, reduced surface shear stress and wake recovery are compared to data for the classical case of a solid backward-facing step. Results provide new insights into the interpretation for flux estimation by eddy-covariance and flux gradient methods and for the assessment of surface boundary conditions in turbulence models of the atmospheric boundary layer in complex landscapes and over water bodies affected by canopy wakes. The wind tunnel results indicate that the wake of a forest canopy strongly affects surface momentum flux within a distance of 35–100 times the step or canopy height, and mean turbulence quantities require distances of at least 100 times the canopy height to adjust to the new surface. The near-surface mixing length in the wake exhibits characteristic length scales of canopy flows at the canopy edge, of the flow separation in the near wake and adjusts to surface layer scaling in the far wake. Components of the momentum budget are examined individually to determine the impact of the canopy wake. The results demonstrate why a constant flux layer does not form until far downwind in the wake. An empirical model for surface shear stress distribution from a forest canopy to a clearing or lake is proposed.     

不同土地覆被下岩溶表层系统CO2体积分数研究

对重庆金佛山林地、裸地表层岩溶生态系统CO2体积分数进行了野外监测,揭示了CO2体积分数变化规律,这种变化与土壤温度有密切的关系。林地与裸地各个层次土壤的CO2体积分数与土温呈一致性变化,随着土温的升高或降低而相应的增加或减少。文章进一步揭示了林地植被平抑这种动态效应,而裸地则响应于温度变化;这种不同植被系统下的动态差异在解释岩溶沉积记录和讨论岩溶作用与碳循环的关系时值得充分注意。     

淮河流域农田碳收支特征及影响因素分析

采用开路涡度相关系统对淮河流域农田湍流特征进行了4年连续监测,以该地区主要农作物小麦和水稻为例,对CO2通量数据进行分析处理。研究结果表明：农田CO2通量的日变化随季节变化明显,春、夏变化幅度明显大于秋、冬。2007—2010年淮河流域夏季农田CO2通量日变化规律均呈单峰型,白天为明显的碳汇;其日最大累计吸收量出现在2007年,可达16.1mg.m-2.s-1,最小值出现在2009年,为11.1 mg.m-2.s-1。在水稻拔节期垂直风速（W）、潜热通量（LE）、显热通量（H）、光合有效辐射（PAR）、净辐射（Rn）与CO2通量均呈负相关,通过对各项指标进行主成分分析,得到了新的综合指标F=-0.36ZX1＋0.42ZX2＋0.42ZX3＋0.42ZX4＋0.39ZX5＋0.42ZX6＋0.13ZX7,并经过计算得出中午1点是最大的碳汇,凌晨12点是最大的碳源。     

Nocturnal accumulation of CO2 underneath a tropical forest canopy along a topographical gradient

Flux measurements of carbon dioxide and water vapor above tropical rain forests are often difficult to interpret because the terrain is usually complex. This complexity induces heterogeneity in the surface but also affects lateral movement of carbon dioxide (CO2) not readily detected by the eddy covariance systems. This study describes such variability using measurements of CO2 along vertical profiles and along a toposequence in a tropical rain forest near Manaus, Brazil. Seasonal and diurnal variation was recorded, with atmospheric CO2 concentration maxima around dawn, generally higher CO2 build-up in the dry season and stronger daytime CO2 drawdown in the wet season. This variation was reflected all along the toposequence, but the slope and valley bottom accumulated clearly more CO2 than the plateaus, depending on atmospheric stability. Particularly during stable nights, accumulation was along lines of equal altitude, suggesting that large amounts of CO2 are stored in the valleys of the landscape. Flushing of this store only occurs during mid-morning, when stored CO2 may well be partly transported back to the plateaus. It is clear that, for proper interpretation of tower fluxes in such complex and actively respiring terrain, the horizontal variability of storage needs to be taken into account not only during the night but also during the mornings.     

珠江三角洲四种森林类型土壤CO2通量特征研究

采用开路式土壤CO2通量测量系统Li-8100&Li-8150对珠江三角洲地区尾叶桉(Eucalyptus urophylla)人工林、乡土树种恢复林、针阔叶混交林和常绿阔叶林4种林型的土壤CO2通量进行了观测。结果表明:4种森林类型年均土壤CO2通量为尾叶桉人工林(3.35μmol.m-2.s-1)>针阔叶混交林(2.66μmol.m-2.s-1)>乡土树种恢复林(2.09μmol.m-2.s-1)>常绿阔叶林(1.86μmol.m-2.s-1);旱季土壤CO2通量明显小于雨季。前3种森林类型凋落物呼吸处理表明,旱季对照组土壤CO2通量均小于相应的去除凋落物组、雨季则相反,全年的对比结果显示,3种森林类型的凋落物呼吸贡献分别达到1.3%、7.1%和10.8%。土壤CO2通量与10 cm土壤温度呈显著指数相关,且土壤CO2通量温度敏感指数表现为针阔叶混交林Q10最大(3.49),尾叶桉人工林Q10最小(1.95)。     

珠江三角洲四种森林类型土壤CO_2通量特征研究

采用开路式土壤CO2通量测量系统Li-8100＆Li-8150对珠江三角洲地区尾叶桉（Eucalyptus urophylla）人工林、乡土树种恢复林、针阔叶混交林和常绿阔叶林4种林型的土壤CO2通量进行了观测。结果表明：4种森林类型年均土壤CO2通量为尾叶桉人工林（3.35μmol.m-2.s-1）〉针阔叶混交林（2.66μmol.m-2.s-1）〉乡土树种恢复林（2.09μmol.m-2.s-1）〉常绿阔叶林（1.86μmol.m-2.s-1）;旱季土壤CO2通量明显小于雨季。前3种森林类型凋落物呼吸处理表明,旱季对照组土壤CO2通量均小于相应的去除凋落物组、雨季则相反,全年的对比结果显示,3种森林类型的凋落物呼吸贡献分别达到1.3%、7.1%和10.8%。土壤CO2通量与10 cm土壤温度呈显著指数相关,且土壤CO2通量温度敏感指数表现为针阔叶混交林Q10最大（3.49）,尾叶桉人工林Q10最小（1.95）。     

Stratified flow interactions with a suspended canopy

Field observations of the interactions between a stratified flow and a canopy suspended from the free surface above a solid boundary are described and analysed. Data were recorded in and around the canopy formed by a large long-line mussel farm. The canopy causes a partial blockage of the water flow, reducing velocities in the upper water column. Deceleration of the approaching flow results in a deepening of isopycnals upstream of the canopy. Energy considerations show that the potential for an approaching stratified flow to be diverted beneath a porous canopy is indicated by a densimetric Froude number. Strong stratification or low-velocities inhibit vertical diversion beneath the canopy, instead favouring a horizontal diversion around the sides. The effect on vertical mixing is also considered with a shear layer generated beneath the canopy and turbulence generated from drag within the canopy. In the observations, stratification is shown to be of sufficient strength to limit the effectiveness of the first mixing process, while the turbulence within the canopy is likely to enhance vertical exchange. Velocity and temperature microstructure measurements are used to investigate the effect of the canopy on turbulent dissipation and show that dissipation is enhanced within the canopy.     

A σ-coordinate transport model coupled with rotational boussinesq-type equations

This paper describes a σ-coordinate scalar transport model coupled with a Boussinesq-type hydrodynamic model. The Boussinesq model has the ability to calculate both three-dimensional velocity distributions and the water surface motion. To capture ‘dispersion’ processes in open channel flow, horizontal vorticity effects induced by a bottom shear stress are included in the Boussinesq model. Thus, a reasonable representation of vertical flow structure can be captured in shallow and wavy flow fields. To solve the coupled Boussinesq and scalar transport system, a finite-volume method, based on a Godunov-type scheme with the HLL Riemann solver, is employed. Basic advection and advection–diffusion numerical tests in a non-rectangular domain were carried out and the computed results show good agreement with analytic solutions. With quantitative comparisons of dispersion experiments in an open channel, it is verified that the proposed coupled model is appropriate for both near and far field scalar transport predictions. From numerical simulations in the surf zone, physically reasonable results showing expected vertical variation are obtained.     

Evidence of large-scale source-sink dynamics and long-distance dispersal among Wood Thrush populations

Source-sink dynamics are commonly thought to occur among Wood Thrush (Hylocichla mustelina) and other songbird populations, allowing for the persistence of populations with negative growth rates ("sinks") through immigration from populations with positive growth rates ("sources"). Knowledge of source-sink dynamics is important for management and conservation because the removal of source habitat should result in the extinction of dependent sinks. However, since research has focused on identifying individual sources/sink populations, not source-sink pairs, we cannot predict these effects or the scale over which they occur. We posit that, when dispersal occurs from a source to a sink year after year, there will be a one-year time-lagged correlation in abundance between the two populations. This should occur for populations separated by distances over which juveniles disperse. Using the North American Breeding Bird Survey data, we tested for such time-lagged correlations between paired Wood Thrush populations from 10 to 200 km apart. Populations were linked with a one-year time lag over distances from 60 to 80 km, indicating that dispersal and source-sink dynamics may occur over these long distances. There was also a declining trend in forest cover from sources to sinks. Conservation and management strategies should therefore be designed at large scales, with consideration for source-sink dynamics and forest cover.     

Tree species control rates of free-living nitrogen fixation in a tropical rain forest

Tropical rain forests represent some of the most diverse ecosystems on earth, yet mechanistic links between tree species identity and ecosystem function in these forests remains poorly understood. Here, using free-living nitrogen (N) fixation as a model, we explore the idea that interspecies variation in canopy nutrient concentrations may drive significant local-scale variation in biogeochemical processes. Biological N fixation is the largest "natural" source of newly available N to terrestrial ecosystems, and estimates suggest the highest such inputs occur in tropical ecosystems. While patterns of and controls over N fixation in these systems remain poorly known, the data we do have suggest that chemical differences among tree species canopies could affect free-living N fixation rates. In a diverse lowland rain forest in Costa Rica, we established a series of vertical, canopy-to-soil profiles for six common canopy tree species, and we measured free-living N fixation rates and multiple aspects of chemistry of live canopy leaves, senesced canopy leaves, bulk leaf litter, and soil for eight individuals of each tree species. Free-living N fixation rates varied significantly among tree species for all four components, and independent of species identity, rates of N fixation ranged by orders of magnitude along the vertical profile. Our data suggest that variations in phosphorus (P) concentration drove a significant fraction of the observed species-specific variation in free-living N fixation rates within each layer of the vertical profile. Furthermore, our data suggest significant links between canopy and forest floor nutrient concentrations; canopy P was correlated with bulk leaf litter P below individual tree crowns. Thus, canopy chemistry may affect a suite of ecosystem processes not only within the canopy itself, but at and beneath the forest floor as well.