Water inputs across a tropical montane landscape in Veracruz,Mexico: synergistic effects of land cover,rain and fog seasonality,and interannual precipitation variability

Land‐cover change can alter the spatiotemporal distribution of water inputs to mountain ecosystems, an important control on land‐surface and land‐atmosphere hydrologic fluxes. In eastern Mexico, we examined the influence of three widespread land‐cover types, montane cloud forest, coffee agroforestry, and cleared areas, on total and net water inputs to soil. Stand structural characteristics, as well as rain, fog, stemflow, and throughfall (water that falls through the canopy) water fluxes were measured across 11 sites during wet and dry seasons from 2005 to 2008. Land‐cover type had a significant effect on annual and seasonal net throughfall (NTF <0=canopy water retention plus canopy evaporation; NTF >0=fog water deposition). Forest canopies retained and/or lost to evaporation (i.e. NTF<0) five‐ to 11‐fold more water than coffee agroforests. Moreover, stemflow was fourfold higher under coffee shade than forest trees. Precipitation seasonality and phenological patterns determined the magnitude of these land‐cover differences, as well as their implications for the hydrologic cycle. Significant negative relationships were found between NTF and tree leaf area index (R²=0.38, P<0.002), NTF and stand basal area (R²=0.664, P<0.002), and stemflow and epiphyte loading (R²=0.414, P<0.001). These findings indicate that leaf and epiphyte surface area reductions associated with forest conversion decrease canopy water retention/evaporation, thereby increasing throughfall and stemflow inputs to soil. Interannual precipitation variability also altered patterns of water redistribution across this landscape. Storms and hurricanes resulted in little difference in forest‐coffee wet season NTF, while El Niño Southern Oscillation was associated with a twofold increase in dry season rain and fog throughfall water deposition. In montane headwater regions, changes in water delivery to canopies and soils may affect infiltration, runoff, and evapotranspiration, with implications for provisioning (e.g. water supply) and regulating (e.g. flood mitigation) ecosystem services.     

热带季节雨林林窗辐射特征研究

利用不同季节热带季节雨林林窗、林内及旷地不同波长太阳辐射的实测资料,探讨了热带季节雨林林窗不同波长辐射特征。结果表明,热带季节雨林林窗中央和北侧林冠下的不同波长太阳辐射存在“突跃现象”,中午前后各辐射量值迅速达到最大后又急速下降;林窗内不同波长太阳辐射日总量值均大于林内,却小于旷地;林窗中央和北侧林冠下比较可见,在干热季、雨季和雨季后期,林窗中央的各辐射要素总量值高于北侧林冠下,但雾凉季时受浓雾和太阳高度以及方位的影响,各辐射要素总量值较北侧林冠下低;林窗中央的总辐射日总量在雨季太阳高度角最大时最高,雨季后期和干热季其次,雾凉季时最小;林窗中央和北侧林冠下的红外辐射及可见光在总辐射中所占份额随季节的变化而不同,充分显示了林窗辐射环境的异质性;与旷地和林内相比,林窗内各测点的红外辐射在总辐射中所占份额介于旷地和林内之间．高于旷地却低于林内;而可见光占总辐射的比值情况正好相反．另外,林窗可见光分配率的季节变化都大于红外辐射;热带季节雨林林窗育增加红外辐射、减少可见光的特征．     

Light use efficiency of California redwood forest understory plants along a moisture gradient

We investigated photosynthesis of five plant species growing in the understory at three sites (1,170-, 1,600- and 2,100-mm annual moisture inputs), along the geographical range of coastal California redwood forest, to determine whether greater inputs of rain and fog at northern sites enhance photosynthetic utilization of fluctuating light. Measurements of understory light environment and gas exchange were carried out to determine steady state and dynamic photosynthetic responses to light. Leaf area index ranged from 4.84 at the 2,100-mm site to 5.98 at the 1,170-mm site. Maximum rates of net photosynthesis and stomatal conductance (g) did not vary appreciably within species across sites. Photosynthetic induction after a change from low to high light was significantly greater in plants growing in lower light conditions regardless of site. Photosynthetic induction also increased with the rate of g in diffuse light, prior to the increase to saturating light levels. Post-illumination CO₂ assimilation was the largest factor contributing to variation in C gain during simulated lightflecks. The duration of post-illumination photosynthetic activity, total CO₂ assimilation per light received, and light use efficiency during simulated lightflecks increased significantly with moisture inputs in four out of five species. Increasing leaf N concentration with increasing moisture inputs in three out of five species, coupled with changes in leaf N isotopic composition with the onset of the summer fog season suggest that natural N deposition increases with rain and fog inputs and contributes to greater utilization of fluctuating light availability in coastal California redwood forests.     

Foliar water uptake: a common water acquisition strategy for plants of the redwood forest

Evaluations of plant water use in ecosystems around the world reveal a shared capacity by many different species to absorb rain, dew, or fog water directly into their leaves or plant crowns. This mode of water uptake provides an important water subsidy that relieves foliar water stress. Our study provides the first comparative evaluation of foliar uptake capacity among the dominant plant taxa from the coast redwood ecosystem of California where crown-wetting events by summertime fog frequently occur during an otherwise drought-prone season. Previous research demonstrated that the dominant overstory tree species, Sequoia sempervirens, takes up fog water by both its roots (via drip from the crown to the soil) and directly through its leaf surfaces. The present study adds to these early findings and shows that 80% of the dominant species from the redwood forest exhibit this foliar uptake water acquisition strategy. The plants studied include canopy trees, understory ferns, and shrubs. Our results also show that foliar uptake provides direct hydration to leaves, increasing leaf water content by 2–11%. In addition, 60% of redwood forest species investigated demonstrate nocturnal stomatal conductance to water vapor. Such findings indicate that even species unable to absorb water directly into their foliage may still receive indirect benefits from nocturnal leaf wetting through suppressed transpiration. For these species, leaf-wetting events enhance the efficacy of nighttime re-equilibration with available soil water and therefore also increase pre-dawn leaf water potentials.     

Foggy days and dry nights determine crown‐level water balance in a seasonal tropical montane cloud forest

The ecophysiology of tropical montane cloud forest (TMCF) trees is influenced by crown‐level microclimate factors including regular mist/fog water inputs, and large variations in evaporative demand, which in turn can significantly impact water balance. We investigated the effect of such microclimatic factors on canopy ecophysiology and branch‐level water balance in the dry season of a seasonal TMCF in Veracruz, Mexico, by quantifying both water inputs (via foliar uptake, FU) and outputs (day‐ and night‐time transpiration, NT). Measurements of sap flow, stomatal conductance, leaf water potential and pressure–volume relations were obtained in Quercus lanceifolia, a canopy‐dominant tree species. Our results indicate that FU occurred 34% of the time and led to the recovery of 9% (24 ± 9.1 L) of all the dry‐season water transpired from individual branches. Capacity for FU was independently verified for seven additional common tree species. NT accounted for approximately 17% (46 L) of dry‐season water loss. There was a strong correlation between FU and the duration of leaf wetness events (fog and/or rain), as well as between NT and the night‐time vapour pressure deficit. Our results show the clear importance of fog and NT for the canopy water relations of Q. lanceifolia.     

Landscape scale variation in the hydrologic niche of California coast redwood

Topoclimatic diversity within forest landscapes can underlie variation in water availability, which may correspond to patterns in habitat suitability of tree species with differing hydrologic niches. However, the trade-off between the collection of data at a fine grain size over large spatial extents has limited comprehensive analyses of landscape scale variation in habitat suitability. We present a fine scale analysis of the roles of topographic gradients in moisture availability, soil water storage and fog frequency in the spatial pattern of habitat suitability for coast redwood Sequoia sempervirens, at 10 m resolution across 34 800 ha and three landscapes spanning approximately one-third of redwood's latitudinal range. A new 10 m resolution presence-absence map derived from airborne imaging spectroscopy was used to characterize current redwood distributions in three landscapes. Access to belowground moisture was assessed through four measures calculated from a high-resolution LiDAR digital elevation model, soil water storage was mapped from the USDA Soil Survey Geographic Database, and fog frequency was calculated from MODIS cloud cover data. Access to belowground moisture and fog predicted variation in suitability as redwood habitat from 22–75% and predicted redwood presence and absence at 10 m resolution with 63–74% accuracy. Across all three landscapes, redwood density consistently decreased with increasing interpolated height above a stream (IHAS), but the role of other predictors varied among the sites. Our results challenge previous assumptions that redwood habitat suitability within much coarser resolutions (800 m × 800 m) can be described by a single value. The elevated habitat suitability of sites close to streams suggests that these sites should be evaluated for their potential to become microrefugia within landscapes that may become climatically-unsuitable for redwoods at a regional scale.     

西双版纳热带季节雨林内雾特征研究

 利用4年（1999～2002）的雾观测资料,对西双版纳热带季节雨林内雾特征进行了观测研究。结果表明,雾首先形成于最上林冠层,林下雾是由上层雾变浓、下沉而来。夜间,雾形成前,气温高于叶表温;雾形成后,气温则低于叶表温。热带雨林内各季节雾日数和雾日频率均高于无林地。热带雨林内平均全年雾日数可达258 d,其中雾季和干热季共占154 d（59.6%）,而雾季的雾日频率高达90%。雾日数的季节变化与各季节雨量呈明显的负相关。雾季,雾在23∶00左右生成,比干热季、雨季分别提前0.7、2.3 h,而消散时间则分别推迟0.8、2.2 h。雾生成和消散时间呈现出较明显的负相关。雾季雾的持续时间达12.2 h·d-1,比干热季、雨季分别长1.5、4.6 h·d-1。全年雾总持续时间占全年时间的39.7%,而雾季的相应值为50.8%。雾的形成不仅凝结了水汽进入森林（全年89.4 mm）,同时也对森林起到了一定的保温作用,这对热带雨林的生存和发展具有至关重要的作用。     

Edge Effects of Linear Canopy Openings on Tropical Rain Forest Understory Microclimate

We investigated microclimatic edge gradients associated with grassy powerlines, paved highways and perennial creeks in wet tropical forest in northeastern Australia during wet and dry seasons. Photosynthetically active radiation, air temperature and vapor pressure deficit, soil temperature, canopy temperature, soil moisture, and air speed in the rain forest understory were measured during traverses perpendicular to the forest edge. Light intensity was elevated near the edges of powerlines, highways, and creeks, but this effect was strongest for creek edges. Air temperature and vapor pressure deficit were elevated near powerline edges in the dry season and highway edges in both wet and dry seasons but were not elevated near creek edges in either season. In contrast, soil moisture was lowered near creek edges but not near either powerline or highway edges. No edge gradients were detected for air speed. Canopy temperature was elevated near highway edges and lowered near powerline edges in the wet season but no edge gradients in canopy temperature were detected near creek edges in either the wet or the dry season. We suggest that these different edge gradients may be largely the result of differences in the fluxes of latent and sensible heat within each type of linear canopy opening, with periodic flood disturbance assisting by maintaining a more open canopy near creek edges. Our data indicate that the nature of the linear canopy opening is at least as important as the width in determining the nature and severity of microclimatic edge effects, analogous to the "matrix effect" of traditional fragmentation studies.     

Polystichum munitum (Dryopteridaceae) varies geographically in its capacity to absorb fog water by foliar uptake within the redwood forest ecosystem

? Premise of the study: Fog provides a critical water resource to plants around the world. In the redwood forest ecosystem of northern California, plants depend on fog absorbed through foliar uptake to stay hydrated during the rainless summer. In this study, we identified regions within the redwood ecosystem where the fern Polystichum munitum canopy most effectively absorbs fog drip that reaches the forest floor. ? Methods: We measured the foliar uptake capacity of P. munitum fronds at seven sites along 700 km of the redwood forest ecosystem. We quantified the canopy cover of P. munitum at each site and estimated how much water the fern canopy can acquire aboveground through fog interception and absorption. ? Key results: Throughout the ecosystem, nocturnal foliar uptake increased the leaf water content of P. munitum by 7.2%, and we estimated that the P. munitum canopy can absorb 5 ± 3% (mean ± SE) of intercepted fog precipitation. Strikingly, P. munitum had the highest foliar uptake capacity in the center of the ecosystem and may absorb 10% more of the fog its canopy intercepts in this region relative to other regions studied. Conversely, P. munitum had no foliar uptake capacity in the southern end of the ecosystem. ? Conclusions: This study shows the first evidence that foliar uptake varies within species at the landscape scale. Our findings suggest that the P. munitum at the southern tip of the redwood ecosystem may suffer most from low summertime water availability because it had no potential to acquire fog as an aboveground water subsidy.     

热带季节雨林林窗边缘不同热力作用面热力效应的季节变化特征

通过对热带季节雨林雾凉季和湿热季昼间林窗区域不同热力作用面的热力效应初步分析,指出在西双版纳,不论是雾凉季还是湿热季,热带季节雨林林窗边缘壁面均具有不可忽视的热力作用,且由于受林缘树木的影响,热力效应较强的东侧,北侧林缘壁面最大区域出现位置高于次生林林窗,而强度小于次生林林窗,显示了林窗边缘壁面的热力效应除与太阳高度角,太阳辐射的时间长短和强度有关之外,林窗边缘树木高度也是不可忽视的因子,其结果可为进一步研究林窗小气候形成机制提供研究基础,为研究林窗更新及生物多样性问题提供科学参考。     

西双版纳热带季节雨林水热通量

利用西双版纳热带季节雨林2003和2004年常规气象、生物量以及水热通量观测资料,对该林地两年内各能量分量的数值大小和变化规律、能量分配以及水量平衡特征等进行了分析研究。结果表明,2003和2004年净辐射总量分别为3516.4MJ/(m.2a)和3516.6MJ/(m.2a)。在能量分配过程中潜热通量占优势,2003年和2004年的总量分别是相应年份净辐射总量的46%和44%,显热通量则分别只有12%和11%。2003年和2004年林冠传导率均值分别为10.3mm/s和10.0mm/s,其中干热季期间的林冠传导率明显低于雾凉季和雨季。林冠传导率与叶面积指数和空气饱和水汽压差值之间分别呈极显著的正、负线性相关关系;它基本上不受土壤含水量的影响,只是当长期无雨或雨量很小导致土壤含水量低于0.15m3/m3时,林冠传导率才与土壤含水量间存在极显著的相关关系。西双版纳热带雨林2003和2004年的蒸散量分别是663mm和634mm,受浓雾和林冠传导率的综合影响,该森林生态系统干季蒸散量低于雨季,这是西双版纳热带季节雨林能够在水热极限条件下生存并良好发育的重要原因。     

Fog in the California redwood forest: ecosystem inputs and use by plants

Fog has been viewed as an important source of moisture in many coastal ecosystems, yet its importance for the plants which inhabit these ecosystems is virtually unknown. Here, I report the results of a 3-year investigation of fog inputs and the use of fog water by plants inhabiting the heavily fog inundated coastal redwood (Sequoia sempervirens) forests of northern California. During the study period, 34%, on average, of the annual hydrologic input was from fog drip off the redwood trees themselves (interception input). When trees were absent, the average annual input from fog was only 17%, demonstrating that the trees significantly influence the magnitude of fog water input to the ecosystem. Stable hydrogen and oxygen isotope analyses of water from fog, rain, soil water, and xylem water extracted from the dominant plant species were used to characterize the water sources used by the plants. An isotopic mixing model was employed to then quantify how much fog water each plant used each month during the 3-year study. In summer, when fog was most frequent, ～19% of the water within S.sempervirens, and ～66% of the water within the understory plants came from fog after it had dripped from tree foliage into the soil; for S.sempervirens, this fog water input comprised 13–45% of its annual transpiration. For all plants, there was a significant reliance on fog as a water source, especially in summer when rainfall was absent. Dependence on fog as a moisture source was highest in the year when rainfall was lowest but fog inputs normal. Interestingly, during the mild El Niño year of 1993, when the ratio of rainfall to fog water input was significantly higher and fog inputs were lower, both the proportion and coefficient of variation in how much fog water was used by plants increased. An explanation for this is that while fog inputs were lower than normal in this El Niño year, they came at a time when plant demand for water was highest (summer). Therefore, proportional use of fog water by plants increased. The results presented suggest that fog, as a meteorological factor, plays an important role in the water relations of the plants and in the hydrology of the forest. These results demonstrate the importance of understanding the impacts of climatic factors and their oscillations on the biota. The results have important implications for ecologists, hydrologists, and forest managers interested in fog-inundated ecosystems and the plants which inhabit them.     

The contribution of fog to the water relations of Sequoia sempervirens (D. Don): foliar uptake and prevention of dehydration

Fog is a defining feature of the coastal California redwood forest and fog inputs via canopy drip in summer can constitute 30% or more of the total water input each year. A great deal of occult precipitation (fog and light rain) is retained in redwood canopies, which have some of the largest leaf area indices known (Westman & Whittaker, Journal of Ecology 63, 493–520, 1975). An investigation was carried out to determine whether some fraction of intercepted fog water might be directly absorbed through leaf surfaces and if so, the importance of this to the water relations physiology of coast redwood, Sequoia sempervirens. An array of complimentary techniques were adopted to demonstrate that fog is absorbed directly by S. sempervirens foliage. Xylem sap transport reversed direction during heavy fog, with instantaneous flow rates in the direction of the soil peaking at approximately 5–7% of maximum transpiration rate. Isotopic analyses showed that up to 6% of a leaf's water content could be traced to a previous night's fog deposition, but this amount varied considerably depending on the age and water status of the leaves. Old leaves, which appear most able to absorb fog water were able to absorb distilled water when fully submersed at an average rate of 0.90 mmol m² s^?1, or about 80% of transpiration rates measured at the leaf level in the field. Sequoia sempervirens has poor stomatal control in response to a drying atmosphere, with rates of water loss on very dry nights up to 40% of midday summer values and rates above 10% being extremely common. Owing to this profligate water use behaviour of S. sempervirens, it appears that fog has a greater role in suppressing water loss from leaves, and thereby ameliorating daily water stress, than in providing supplemental water to foliar tissues per se. Although direct foliar absorption from fog inputs represents only a small fraction of the water used each day, fog's in reducing transpiration and rehydrating leaf tissues during the most active growth periods in summer may allow for greater seasonal carbon fixation and thus contribute to the very fast growth rates and great size of this species.     

西双版纳热带雨林干季林冠层雾露形成的小气候特征研究

对西双版纳热带雨林干季林冠层雾露形成的小气候特征进行了观测研究。结果表明,雾露首先形成于最上林冠层,林下露水的形成迟于林上3～4h,林下雾是由上层雾变浓、下沉而来。夜间,雾形成前,气温高于叶表温;雾形成后,气温则低于叶表温,且气温及叶表温均有回升。雾露的形成不仅凝结了水汽进入森林,同时也对森林起到了一定的保温作用,这对热带雨林的生存和发展具有致关重要的作用。     

Regeneration patterns and persistence of the fog-dependent Fray Jorge forest in semiarid Chile during the past two centuries

The persistence of rainforest patches at Fray Jorge National Park (FJNP) in semiarid Chile (30°40′S), a region receiving approximately 147 mm of annual rainfall, has been a source of concern among forest managers. These forests are likely dependent on water inputs from oceanic fog and their persistence seems uncertain in the face of climate change. Here, we assessed tree radial growth and establishment during the last two centuries and their relation to trends in climate and canopy disturbance. Such evaluation is critical to understanding the dynamics of these semiarid ecosystems in response to climate change. We analyzed forest structure of six forest patches (0.2–22 ha) in FJNP based on sampling within 0.1 ha permanent plots. For the main canopy species, the endemic Aextoxicon punctatum (Aextoxicaceae), we used tree‐ring analysis to assess establishment periods, tree ages, growing trends and their relation to El Niño Southern Oscillation (ENSO), rainfall, and disturbance. The population dynamics of A. punctatum can be described by a continuous regeneration mode. Regeneration of A. punctatum was sensitive to different canopy structures. Growth release patterns suggest the absence of large scale human impact. Radial growth and establishment of A. punctatum were weakly correlated with rainfall and ENSO. If water limits forests patch persistence, patches are likely dependent on the combination of fog and rain water inputs. Forest patches have regenerated continuously for at least 250 years, despite large fluctuations in rainfall driven by ENSO and a regional decline in rainfall during the last century. Because of the positive influence on fog interception, forest structure should be preserved under any future climate scenario. Future research in FJNP should prioritize quantifying the long‐term trends of fog water deposition on forests patches. Fog modeling is crucial for understanding the interplay among physical drivers of water inputs under climate change.     

西双版纳热带次生林林窗小气候要素的时空分布特征

利用西双版纳雾凉季和干热季热带次生林林窗的小气候垂直观测资料,探讨了昼间林窗区域树表温、气温、水汽压及相对湿度的时空分布和变化规律,指出在林窗区域,林窗边缘不仅具有显著的热力效应,同样具有明显的水汽效应。并由此构成了林窗区域立体空间的环境异质性,其结果对深入探讨林窗区域的热量、水汽传输,小气候的形成机制,生物多样性和更新等问题均具有重要意义。利用西双版纳雾凉季和干热季热带次生林林窗的小气候垂直观测资料,探讨了昼间林窗区域树表温、气温、水汽压及相对湿度的时空分布和变化规律,指出在林窗区域,林窗边缘不仅具有显著的热力效应,同样具有明显的水汽效应。并由此构成了林窗区域立体空间的环境异质性,其结果对深入探讨林窗区域的热量、水汽传输,小气候的形成机制,生物多样性和更新等问题均具有重要意义。     

Rain Forest Islands in the Chilean Semiarid Region: Fog-dependency, Ecosystem Persistence and Tree Regeneration

Tree presence in semiarid ecosystems is generally constrained by insufficient annual rainfall. However, in semiarid Chile, rainforest patches dominated by Aextoxicon punctatum are unexpectedly found on coastal mountaintops (450–600 m) at 30°S, surrounded by a xerophytic vegetation matrix that receives only 147 mm of annual precipitation. It has been proposed that these forests persist as a result of fog-water inputs. If so, then because fog-water deposition is spatially heterogeneous and shows strong edge effects, the potential environmental gradient created by the direction of fog input should determine forest structure and tree regeneration patterns. To investigate this hypothesis, we measured fog inputs, forest structural attributes (age and size distribution, basal area, and coarse woody debris), and tree regeneration in three different habitats: the windward edges (WE), leeward edges (LE), and the interior (center) of rainforest patches varying in area from 0.2 to 22 ha. Mean fog-water input was estimated from passive collectors over 1 year in WE and LE of patches. Tree regeneration was greater in the WE and forest interior (FI) and decreased toward the LE of patches, following a marked pattern of decline in fog inputs. Older trees and coarse woody debris were concentrated in the FI and LE of patches. Tree regeneration and patch structure appear to be largely controlled by fog-input direction and edge effects. We propose that forest patches may be slowly growing toward the incoming fog edge, while dying at the opposite edge.     

Wet- vs. Dry-Season Transpiration in an Amazonian Rain Forest Palm Iriartea deltoidea

The wet and dry seasons in tropical rain forests can differ in precipitation, soil moisture and irradiance more significantly than often assumed. This could potentially affect the water relations of many tree species that may exhibit either increased transpiration in the dry season as a response to the increased irradiance or decreased transpiration as a result of decreases in soil moisture and increases in atmospheric vapor pressure deficit (VPD). Atmospheric data, soil moisture data and sap fluxes in Iriartea deltoidea palms were measured in eastern Ecuador during the wet and dry seasons. There were no differences between total daily sap fluxes in I. deltoidea palms during the wet and dry seasons; however, evaporative demand was significantly higher in the dry season and therefore, transpiration was more restricted by stomatal closure during the dry season than the wet season. This is likely the result of larger atmospheric VPD during the dry season compared with the wet season and possibly the result of reduced soil moisture availability. Additionally, based on published tree abundances in this area, measured sap fluxes in I. deltoidea were scaled up to the hectare level. Transpiration from I. deltoidea palms was estimated to be around 0.03 mm/d, which could represent about 1 percent of total transpiration in this area of the Amazon rain forest. If climate change predictions for more lengthy tropical dry periods are realized, greater stomatal control of dry-season sap flux has the potential to become even more prevalent in tropical species.     

Spatial and Temporal Variation of Seed Rain in the Canopy and on the Ground of a Tropical Cloud Forest

Spatial and temporal patterns of seed rain impact plant fitness, genetic and demographic structure of plant populations, and species' interactions. Because plants are sessile, they rely on biotic and abiotic dispersal agents to move their seeds. The relative importance of these dispersal agents may shift throughout the year. In tropical forests, seed dispersal of epiphytes constitutes a major but hitherto unknown portion of seed rain ecology. For the first time, we report on patterns of seed rain for both epiphytic and terrestrial plants across an entire year in a Neotropical montane forest. To examine seed rain, we placed traps in the canopy and on the ground. We analyzed seed dispersal syndrome (bird, mammal, wind) and plant habit (epiphyte, liana, shrub, small tree, large tree) across all seasons of the year (dry, misty, wet). We found that the community of species collected in canopy traps was significantly different from the community in ground traps. Epiphytes were the most common plant habit found in canopy traps, while large trees were most common in ground traps. Species with bird‐dispersed seeds dominated all traps. Species richness was significantly higher during the dry season in ground traps, but did not vary across seasons in canopy traps. Our results highlight the distinct seed rain found in the canopy and on the ground and underscore the importance of frugivores for dispersing both arboreal and terrestrial plants in tropical ecosystems.     

Large Canopy Exchange Fluxes of Inorganic and Organic Nitrogen and Preferential Retention of Nitrogen by Epiphytes in a Tropical Lowland Rainforest

Little is known about how tropical forest canopies interact with atmospheric nitrogen deposition and how this affects the internal nutrient dynamics and the processing of external nutrient inputs. The objectives of this study therefore were (1) to investigate gross and net canopy nitrogen (N) fluxes (retention and leaching) and (2) the effect of canopy components on net canopy N retention. Tracers were applied on detached branches in a tropical wet lowland rainforest, Costa Rica. A novel ¹⁵N pool dilution method showed that gross canopy fluxes (retention and leaching) of NO₃ ^?, NH₄ ⁺, and dissolved organic nitrogen (DON) were remarkably higher than net throughfall fluxes. Gross fluxes of NH₄ ⁺ and NO₃ ^? resulted in a negligible net flux whereas DON showed net uptake by the canopy. The highest quantity of ¹⁵N was recovered in epiphytic bryophytes (16.4%) although the largest biomass fraction was made up of leaves. The study demonstrates that tracer applications allow investigation of the dynamic and complex canopy exchange processes and that epiphytic communities play a major role in solute fluxes in tree canopies and therefore in the nutrient dynamics of tropical rain forests.