Complex consequences of disturbance on canopy plant communities of world forests: a review and synthesis.

Epiphytes and their associated biota are increasingly recognized as contributing to biodiversity and to filling critical ecosystem functions in world forests. However, the attributes that have made them successful in canopy environments also make them vulnerable to natural and human-induced disturbances. Drawing upon ecological frameworks to understand disturbance, I categorized and synthesized the drivers and the consequences of disturbances on epiphytic materials. Across all impacts, disturbance agents were significantly more likely to lead to negative, rather than positive, effects in both tropical and temperate locales. Significantly more studies reported negative effects on abundance, diversity, community composition and connectivity, but some studies showed that disturbances enhanced these attributes. Although particular disturbance agents did not differently influence individual consequences, they explained a significant portion of variation in aggregated totals. Surprisingly, relative to human disturbances, natural disturbances were more likely to lead to negative effects. Many studies provided recommendations for effective societal responses to mitigate negative impacts, such as retaining large, old trees in forestry operations, patch-clearing for epiphyte harvest, maximizing forest fragment size, using epiphytes as bioindicators of disturbance, and applying principles of community forestry to land management. Future actions should also include communication of these results to policymakers and land managers.

Variation in cloud immersion, not precipitation, drives leaf trait plasticity and water relations in vascular epiphytes during an extreme drought.

PREMISE: Epiphytes are abundant in ecosystems such as tropical montane cloud forests where low-lying clouds are often in contact with vegetation. Climate projections for these regions include more variability in rainfall and an increase in cloud base heights, which would lead to drier conditions in the soil and atmosphere. While recent studies have examined the effects of drought on epiphytic water relations, the influence that atmospheric moisture has, either alone or in combination with drought, on the health and performance of epiphyte communities remains unclear. METHODS: We conducted a 10-week drought experiment on seven vascular epiphyte species in two shadehouses, one with warmer and drier conditions and another that was cooler and more humid. We measured water relations across control and drought-treatment groups and assessed functional traits of leaves produced during drought conditions to evaluate trait plasticity. RESULTS: Epiphytes exposed to drought and drier atmospheric conditions had a significant reduction in stomatal conductance and leaf water potential and an increase in leaf dry matter. Nonsucculent epiphytes from the drier shadehouse had the greatest shifts in functional traits, whereas succulent epiphytes released stored leaf water to maintain water status. CONCLUSIONS: Individuals in the drier shadehouse had a substantial reduction in performance, whereas drought-treated individuals that experienced cloud immersion displayed minimal changes in water status. Our results indicate that projected increases in the cloud base height will reduce growth and performance of epiphytic communities and that nonsucculent epiphytes may be particularly vulnerable.

Gigantic chloroplasts, including bizonoplasts, are common in shade-adapted species of the ancient vascular plant family Selaginellaceae.

PREMISE: Unique among vascular plants, some species of Selaginella have single giant chloroplasts in their epidermal or upper mesophyll cells (monoplastidy, M), varying in structure between species. Structural variants include several forms of bizonoplast with unique dimorphic ultrastructure. Better understanding of these structural variants, their prevalence, environmental correlates and phylogenetic association, has the potential to shed new light on chloroplast biology unavailable from any other plant group. METHODS: The chloroplast ultrastructure of 76 Selaginella species was studied with various microscopic techniques. Environmental data for selected species and subgeneric relationships were compared against chloroplast traits. RESULTS: We delineated five chloroplast categories: ME (monoplastidy in a dorsal epidermal cell), MM (monoplastidy in a mesophyll cell), OL (oligoplastidy), Mu (multiplastidy, present in the most basal species), and RC (reduced or vestigial chloroplasts). Of 44 ME species, 11 have bizonoplasts, cup-shaped (concave upper zone) or bilobed (basal hinge, a new discovery), with upper zones of parallel thylakoid membranes varying subtly between species. Monoplastidy, found in 49 species, is strongly shade associated. Bizonoplasts are only known in deep-shade species (<2.1% full sunlight) of subgenus Stachygynandrum but in both the Old and New Worlds. CONCLUSIONS: Multiplastidic chloroplasts are most likely basal, implying that monoplastidy and bizonoplasts are derived traits, with monoplastidy evolving at least twice, potentially as an adaptation to low light. Although there is insufficient information to understand the adaptive significance of the numerous structural variants, they are unmatched in the vascular plants, suggesting unusual evolutionary flexibility in this ancient plant genus.

Contrasting effects of host tree isolation on population connectedness in two tropical epiphytic bromeliads.

PREMISE: Conversion of primary forests to pastures is a major cause of habitat fragmentation in the tropics. Fragmentation is expected to impede gene flow for many plant species that are restricted to remaining forest fragments. Epiphytes may be especially vulnerable to this effect of forest fragmentation because they depend on host trees. However, trees that remain in pastures may enhance connectivity across the landscape for epiphyte species that can thrive on such trees. To investigate this possibility, we studied the genetic structures of two such species on isolated pasture trees and surrounding forest, in relation to their local abundances in different habitat types and aspects of their reproductive biology including pollen and seed dispersal agents, and looked for evidence of increased or diminished gene flow. METHODS: We used microsatellite markers to assess geographic patterns of genetic diversity and differentiation in two epiphytic bromeliads, Catopsis nitida and Werauhia tonduziana, in the Monteverde region of Costa Rica. RESULTS: About 85% of the FST value for Catopsis nitida was found among pastures within regions, while for Weruahia tonduziana, about 80% of the FST value was contributed by differences between regions, indicating much more gene flow within regions, relative to C. nitida. CONCLUSIONS: Although there was substantial genetic differentiation among epiphyte populations, those on isolated pasture trees were not substantially less diverse than those in adjacent forests, suggesting that pasture trees may serve as "stepping stones" that help these species maintain their genetic connectedness and diversity at larger geographic scales.

After the Fall: The Tapestry of Disturbance and Recovery.

On July 3, 2015, Nalini Nadkarni, a world-renowned ecologist who had been studying the biologic processes of ecosystem disturbance and recovery, sustained a catastrophic 50-foot free-fall from the top of the rainforest canopy to the forest floor at her remote field research site. She lost consciousness in shock and sustained life-threatening injuries. Her accompanying students hiked out, radio-called 911, and the Harborview Medical Center (Seattle) Medivac team arrived 4 hours later to rescue her. Her prognosis was extremely grim; her family gathered in anticipation of her death as she underwent four operations during her 10 days in the ICU. As she emerged from coma, she spent weeks of hospital recovery and months of progressive mobility and physical therapy during medical leave from work. She experienced ICU psychosis and postintensive care syndrome, but slowly recovered nearly totally, to the point where she can solo hike up to 18 miles in a day, and has fully resumed her professional responsibilities as professor of biology, including climbing tall trees for her canopy research. She attributes her survival and remarkable recovery to both exquisite medical critical care and support she received, and also to incorporating lessons learned from her interdisciplinary study of how diverse natural systems commonly experience and recover from catastrophic disturbances (e.g., forest fires, traffic jams, orphaned children, and refugee survivors of war). Insights from her own encounter with critical illness and study of disturbance and recovery led her to reflect on the tapestry of disturbance and recovery that permeate all ecosystems, and with relevance to the evolving Society of Critical Care Medicine, postintensive care syndrome, and THRIVE initiatives.

Providing Public Engagement Training to Build Connections Between the Community and Research Ethics Professionals: A Pilot Project.

There is growing interest for research ethics professionals to engage with members of the public, yet they often lack the training needed to engage effectively. The STEM Ambassador Program provides a promising framework for training research ethics professionals to form authentic community connections and carry out effective engagement activities based on shared interests and values. The experiences of ten research administrators who participated in a pilot of the STEM Ambassador training for research ethics professionals are presented. Post-training surveys of the research administrators indicate that they valued the training and the skills obtained, and intend to continue with public engagement activities with support of their leadership.

The Future of Critical Care: Optimizing Technologies and a Learning Healthcare System to Potentiate a More Humanistic Approach to Critical Care.

While technological innovations are the invariable crux of speculation about the future of critical care, they cannot replace the clinician at the bedside. This article summarizes the work of the Society of Critical Care Medicine-appointed multiprofessional task for the Future of Critical Care. The Task Force notes that critical care practice will be transformed by novel technologies, integration of artificial intelligence decision support algorithms, and advances in seamless data operationalization across diverse healthcare systems and geographic regions and within federated datasets. Yet, new technologies will be relevant and meaningful only if they improve the very human endeavor of caring for someone who is critically ill.

Frankia and Alnus rubra canopy roots: an assessment of genetic diversity, propagule availability, and effects on soil nitrogen.

The ecological importance of microbial symbioses in terrestrial soils is widely recognized, but their role in soils that accumulate in forest canopies is almost entirely unknown. To address this gap, this study investigated the Frankia-Alnus rubra symbiosis in canopy and forest floor roots at Olympic National Park, WA, USA. Sixteen mature A. rubra trees were surveyed and Frankia genetic diversity in canopy and forest floor nodules was assessed with sequence-based nifH analyses. A seedling bioassay experiment was conducted to determine Frankia propagule availability in canopy and forest floor soils. Total soil nitrogen from both environments was also quantified. Nodules were present in the canopies of nine of the 16 trees sampled. Across the study area, Frankia canopy and forest floor assemblages were similar, with both habitats containing the same two genotypes. The composition of forest floor and canopy genotypes on the same tree was not always identical, however, suggesting that dispersal was not a strictly local phenomenon. Frankia seedling colonization was similar in canopy soils regardless of the presence of nodules as well as in forest floor soils, indicating that dispersal was not likely to be a major limiting factor. The total soil nitrogen of canopy soils was higher than that of forest floor soils, but the presence of Frankia nodules in canopy soils did not significantly alter soil nitrogen levels. Overall, this study indicates that the Frankia-A. rubra symbiosis is similar in canopy and forest floor environments. Because canopy roots are exposed to different environmental conditions within very small spatial areas and because those areas can be easily manipulated (e.g., fertilizer or watering treatments), they present microbial ecologists with a unique arena to examine root-microbe interactions.

How repeated exposure to informal science education affects content knowledge of and perspectives on science among incarcerated adults.

Over two million men, women, and youth are incarcerated in the United States. This large and ethnically diverse population has, in general, more limited exposure to education, particularly in scientific fields, than the general public. Formal educational programs for the incarcerated can be expensive and logistically difficult to initiate and maintain, but informal science education (ISE) approaches have the potential to significantly improve inmates' view of science and of themselves as science learners. However, "dosage effects"-how repeated exposure to educational experiences may affect listeners-are poorly documented. In this study, we evaluated the longitudinal effects of an ISE program in Utah, which provided a monthly lecture series delivered by academic scientists on a range of science topics. Science content knowledge, self-perception as a science learner, interest in science, and a desire to seek out more scientific information all significantly improved for inmates attending lectures. We also found that seeing a greater number of lectures is positively associated with a desire to seek out additional information. We documented an inverse relationship between education background and the increase in a desire to learn more, suggesting that those with more limited exposure to science manifest the greatest increase in seeking out more information. These results suggest that ISE for the incarcerated significantly improves their knowledge of, and relationship with, science; that some of these effects carry over across months or years; and that ISE programs can have the largest impact by focusing on those with more limited prior exposure to science.

Effects of Astrobiology Lectures on Knowledge and Attitudes about Science in Incarcerated Populations.

The incarcerated population has little or no access to science education programs, STEM resources, or scientists. We explored the effects of a low-cost, potentially high-impact informal science education program that enabled NASA scientists to provide astrobiology lectures to adults inside 16 correctional institutions in three states. Post- versus pre-lecture surveys suggest that presentations significantly increased science content knowledge, positively shifted attitudes about science and scientists, increased a sense of science self-identity, and enhanced behavioral intentions about communicating science. These were significant across ethnicity, gender, education level, and institution type, size, location, and state. Men scored higher than women on pre-lecture survey questions. Among men, participants with greater levels of education and White non-Hispanics scored higher than those with less educational attainment and African American and other minority participants. Increases in science content knowledge were greater for women than men and, among men, for those with lower levels of education and African American participants. Women increased more in science identity than did men. Thus, even limited exposure to voluntary, non-credit science lectures delivered by scientists can be an effective way to broker a relationship to science for this underserved public group and can potentially serve as a step to broaden participation in science.

Canopy seed banks as time capsules of biodiversity in pasture-remnant tree crowns.

Tropical pastures present multiple barriers to tree regeneration and restoration. Relict trees serve as "regeneration foci" because they ameliorate the soil microclimate and serve as safe spots for dispersers. Here, we describe another mechanism by which remnant trees may facilitate pasture regeneration: the presence of seed banks in the canopy soil that accumulates from decomposing epiphytes within the crowns of mature remnant trees in tropical cloud forest pastures. We compared seed banks of canopy soils (histosols derived from fallen leaves, fruits, flower, and twigs of host trees and epiphytes, dead bryophytes, bark, detritus, dead animals, and microorganisms, and dust that accumulate on trunks and the upper surfaces of large branches) in pastures, canopy soils in primary forest trees, and soil on the forest floor in Monteverde, Costa Rica. There were 5211 epiphytic and terrestrial plant seeds in the three habitats. All habitats were dominated by seeds in a relatively small number of plant families, most of which were primarily woody, animal pollinated, and animal dispersed. The density of seeds on the forest floor was greater than seed density in either pasture-canopy or forest-canopy soils; the latter two did not differ. Eight species in 44 families and 61 genera from all of the habitats were tallied. There were 37 species in the pasture-canopy soil, 33 in the forest-canopy soil, and 57 on the forest floor. Eleven species were common to all habitats. The mean species richness in the pasture canopy was significantly higher than the forest canopy (F =83.38; p < 0.02). Nonmetric multidimensional scaling ordination revealed that the communities were distinct. Greenhouse experiments verified that many of these seeds were viable, with 29 taxa germinating (23 taxa in pruned mats [mimic of exposed conditions] and 16 taxa in control mats [intact conditions]) within 2 months of observation. Nearly half the species that germinated were characteristic of primary forests (primary forest samples, 19%; pasture samples, 29%). This supports the idea that canopy seed banks of pasture trees can function as time capsules by providing propagules that are removed in both space and time from the primary forest. Their presence may enhance the ability of pastures to regenerate more quickly, reinforcing the importance of trees in agricultural settings.

Canopy soil bacterial communities altered by severing host tree limbs.

Trees of temperate rainforests host a large biomass of epiphytic plants, which are associated with soils formed in the forest canopy. Falling of epiphytic material results in the transfer of carbon and nutrients from the canopy to the forest floor. This study provides the first characterization of bacterial communities in canopy soils enabled by high-depth environmental sequencing of 16S rRNA genes. Canopy soil included many of the same major taxonomic groups of Bacteria that are also found in ground soil, but canopy bacterial communities were lower in diversity and contained different operational taxonomic units. A field experiment was conducted with epiphytic material from six Acer macrophyllum trees in Olympic National Park, Washington, USA to document changes in the bacterial communities of soils associated with epiphytic material that falls to the forest floor. Bacterial diversity and composition of canopy soil was highly similar, but not identical, to adjacent ground soil two years after transfer to the forest floor, indicating that canopy bacteria are almost, but not completely, replaced by ground soil bacteria. Furthermore, soil associated with epiphytic material on branches that were severed from the host tree and suspended in the canopy contained altered bacterial communities that were distinct from those in canopy material moved to the forest floor. Therefore, the unique nature of canopy soil bacteria is determined in part by the host tree and not only by the physical environmental conditions associated with the canopy. Connection to the living tree appears to be a key feature of the canopy habitat. These results represent an initial survey of bacterial diversity of the canopy and provide a foundation upon which future studies can more fully investigate the ecological and evolutionary dynamics of these communities.

Factors affecting the initiation and growth of aboveground adventitious roots in a tropical cloud forest tree: an experimental approach.

Some of the proximate factors that would induce aboveground stems to produce adventitious roots were investigated experimentally on Senecio cooperi, a tropical cloud forest tree. Stem segments were air-layered with different treatments to promote root formation, and the number of roots initiated and rates of root growth were monitored for 20 weeks. Treatments were the application of wet epiphytes or dry epiphytes plus associated humus, sponges wetted with either water or nutrient solutions, or dry sponges. Controls (stem segments with nothing applied) were also monitored. Numbers of adventitious roots formed and rates of subsequent root growth differed among treatments. Wet epiphyte/humus and nutrient solutions were most effective in producing roots, which suggests that epiphytes and the nutrients they intercept and retain within the canopy may cue adjacent host tissue to exploit this resource.

Potential effects of climate change on canopy communities in a tropical cloud forest: an experimental approach.

Global climate change models predict reduced cloud water in tropical montane forests. To test the effects of reduced cloud water on epiphytes, plants that are tightly coupled to atmospheric inputs, we transplanted epiphytes and their arboreal soil from upper cloud forest trees to trees at slightly lower elevations that are naturally exposed to less cloud water. Control plants moved between trees within the upper site showed no transplantation effects, but experimental plants at lower sites had significantly higher leaf mortality, lower leaf production, and reduced longevity. After the epiphytes died, seedlings of terrestrial gap-colonizing tree species grew from the seed banks within the residual mats of arboreal soil. Greenhouse experiments confirmed that the death of epiphytes can result in radical compositional changes of canopy communities. Thus, tropical montane epiphyte communities constitute both a potentially powerful tool for detecting climate changes and a rich arena to study plant/soil/seed interactions under natural and manipulated conditions. This study also provides experimental evidence that the potential effects of global climate change on canopy and terrestrial communities can be significant for cloud forest biota. Results suggest there will be negative effects on the productivity and longevity of particular epiphytes and a subsequent emergence of an emerging terrestrial component into the canopy community from a previously suppressed seed bank.

Nitrogen-15 natural abundance in a montane cloud forest canopy as an indicator of nitrogen cycling and epiphyte nutrition.

Nutrients obtained by epiphytes may either be of atmospheric origin or from organic matter in the canopy, which decomposes to form canopy soil on large branches. We hypothesised that the N supply for epiphytes on small branches was lower, and a larger proportion provided by rainwater, than for epiphytes rooting in canopy soil. We tested this by measuring the N concentration and isotopic composition in terrestrial and canopy soil and in various canopy compartments of a Costa Rican cloud forest. In general, epiphytes on small branches without canopy soil had lower N foliar concentrations and δ15N signals than plants rooted in canopy soil, suggesting that the former receive a higher proportion of N directly from the rain. Epiphytes on small branches also had less negative δ13C values, indicating more frequent water stress. Epiphytes had lower δ15N values (-3.9±2.3‰) than ground-rooted trees (-1.1±1.6‰), and canopy soil had lower values (0.7±1.2‰) than terrestrial soil (3.8±0.7‰). Assuming that the isotopic effect of terrestrial and canopy soil organic matter formation is similar, our findings support earlier results showing that canopy soil is derived mainly from epiphytes, with only minor inputs from host tree matter. Thus, the epiphyte N cycle appears to be largely detached from the tree-soil cycle. Epiphylls on leaves of understorey shrubs had higher δ15N signals than cryptogams in the upper canopy, as a result of either 15N accumulation in throughfall or increased N2 fixation. The correlation between epiphyll and understorey host leaf δ15N suggests some exchange of N between epiphylls and host leaves. Differences between epiphyte groups also appear to be related to uptake of N through mycorrhizas or N2 fixation. Thus, the source and quantity of N supply is highly variable, depending on the systematic group and canopy position.

Reproductive phenology of epiphytes in Monteverde, Costa Rica

Phenology of plants, or the timing of life cycle events, is important for understanding plant ecology, forest dynamics, and plant-animal interactions. In tropical forests, studies that document epiphyte reproductive phenology are relatively few because of the challenges of tracking plants that live in the canopy. Phenological patterns for 279 individuals of 7 epiphyte species were examined across 12 months in a tropical montane forest in Monteverde, Costa Rica. Epiphytes were located in one of two common tree species, Ficus tuerckheimii (Moraceae) or Ocotea tonduzii (Lauraceae). Flowering and fruiting (i.e., when ripe or unripe fruit is present on the plant) of study plants was recorded on monthly intervals, and phenology was examined as a function of the season at the study site (i.e., wet, transition, or dry season), and pollinator syndrome (bird-, or insect-pollinated) and seed dispersal syndrome (bird-, bat-, or wind-dispersed) of each plant. Though some epiphyte species flowered and fruited throughout the year, the majority showed significant seasonality in phenological events. Based on circular statistics, the timing of mean flowering of different epiphyte species varied, however, timing of mean fruiting for most species tended to occur during the wet season. Insect- and bird-pollinated species had peak flowering during the dry season and late wet season, respectively. Bird-dispersed fruits were present each month of the year with peaks from February to October and again in December. Wind-dispersed fruits were observed eight months of the year with a peak in the early wet season. The timing of epiphyte flowering coincided with flowering of large trees in the area. Epiphyte fruiting, however, is distinct from large tree fruiting. Our results demonstrate the seasonal nature of flowering and fruiting in individual epiphyte species while also highlighted the asynchronous nature of phenological events amongst the epiphyte community.

La fenología de las plantas, o el cronograma de eventos en el ciclo de vida, es importante para la comprensión de la ecología vegetal, la dinámica de los bosques y de las interacciones planta-animal. En los bosques tropicales, los estudios que documentan la fenología de las epífitas son relativamente pocos debido a los desafíos que representa darles seguimiento a las plantas que viven en el dosel. Aquí se presenta un reporte de los patrones fenológicos de 279 individuos de 7 especies de epífitas que abarca 12 meses en un bosque montano tropical en Monteverde, Costa Rica. Las epífitas se encuentran en una de las dos especies de árboles comunes, Ficus tuerckheimii (Moraceae) u Ocotea tonduzii (Lauraceae). Se registró la floración y fructificación (i.e. cuando la fruta madura o inmadura está presente en la planta) de las plantas de estudio en intervalos mensuales y examinó la fenología a través de las estaciones en el sitio de estudio (i.e. estación húmeda, transición, o seca) y el síndrome de polinizadores (ave- , o insecto-polinización ) y el síndrome de dispersión de semillas (ave-, murciélago-, o viento- dispersión) de cada planta. Aunque la mayoría de las especies de epífitas tuvieron flores y frutos durante todo el año, la mayoría mostró estacionalidad significativa en los eventos fenoló-gicos. Con base en estadísticas circulares, el tiempo de floración promedio de las especies de epífitas es variado, sin embargo, el momento de la fructificación promedio para la mayoría de las especies tiende a ocurrir durante la estación húmeda. Especies de insectos y aves de polinización tenían pico de floración durante la estación seca y la estación lluviosa tarde, respectivamente. Frutas dispersadas por aves estaban presentes todos los meses del año con picos de febrero a octubre y de nuevo en diciembre. Frutas dispersadas por el viento se observaron ocho meses del año con un pico en la temporada de lluvias temprana. El momento de la floración de las epífitas coincidió con la floración de árboles de gran tamaño en la zona. La fructificación de las epífitas, sin embargo, fue diferente de la fructificación de los árboles de gran tamaño. Nuestros resultados ponen de manifiesto el carácter estacional de la floración y fructificación de las especies epífitas individuales, además de destacar el carácter asincrónico de los eventos fenológicos entre toda la comunidad de epífitas.

Epiphytic and terrestrial mycorrhizas in a lower montane Costa Rican cloud forest.

The epiphyte community is the most diverse plant community in neotropical cloud forests and its collective biomass can exceed that of the terrestrial shrubs and herbs. However, little is known about the role of mycorrhizas in this community. We assessed the mycorrhizal status of epiphytic (Araceae, Clusiaceae, Ericaceae, and Piperaceae) and terrestrial (Clusiaceae, Ericaceae) plants in a lower montane cloud forest in Costa Rica. Arbuscular mycorrhizas were observed in taxa from Araceae and Clusiaceae; ericoid mycorrhizas were observed in ericaceous plants. This is the first report of intracellular hyphal coils characteristic of ericoid mycorrhizas in roots of Cavendishia melastomoides, Disterigma humboldtii, and Gaultheria erecta. Ericaceous roots were also covered by an intermittent hyphal mantle that penetrated between epidermal cells. Mantles, observed uniquely on ericaceous roots, were more abundant on terrestrial than on epiphytic roots. Mantle abundance was negatively correlated with gravimetric soil water content for epiphytic samples. Dark septate endophytic (DSE) fungi colonized roots of all four families. For the common epiphyte D. humboldtii, DSE structures were most abundant on samples collected from exposed microsites in the canopy. The presence of mycorrhizas in all epiphytes except Peperomia sp. suggests that inoculum levels and environmental conditions in the canopy of tropical cloud forests are generally conducive to the formation of mycorrhizas. These may impact nutrient and water dynamics in arboreal ecosystems.

Nitrogen isotope ratios shift with plant size in tropical bromeliads.

We describe an ontogenetic shift in nitrogen (N) isotopic values in two rosette-forming epiphytic bromeliads. Leaf tissue N isotope values of small individuals of two bromeliad species (mean -6.2 per thousand ) differed from those of large individuals within each species (mean -0.5 per thousand ). Using references for potential N sources, we calculated the relative contribution of autochthonous (soil-derived through leaf litter) and allochthonous (atmospheric deposition) N with a two-member mixing model. Atmospheric sources contributed as much as 77-80% of the N in small individuals, whereas soil-derived N contributed 64-72% (conservative reference value) to 100% (less conservative reference value) of leaf tissue N in large plants. Shifts in N source with increasing plant size may be important aspects of rainforest complexity, an understudied aspect of ecosystem diversity.