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Cloud forests have been found to lose more nitrogen in stream discharge than they gain from atmospheric deposition. They also support a large diversity and biomass of tree epiphytes, predominately composed of cryptogams. Since cryptogam epiphytes harbor nitrogen fixing cyanobacteria, they may help make up for the nitrogen loss from ecosystems. We assessed cryptogam biomass on the ground, boles and branches in Quercus costaricensis dominated stands near the tree line in the Cordillera de Talamanca, Costa Rica. Nitrogen fixation was assayed using 15N2 uptake. Total cryptogam biomass was 2 977 kg ha−1, with 67% being found on the lower branches. Bryophytes and chlorolichens made up 53% and 44%, respec- tively, of the biomass. Half of the bryophyte mass was composed of the liverwort Plagiochila heterophylla, and 66% of the chlorolichen of Lobariella pallida. There were no significant differences in nitrogen fixation rates between the cryptogam species, with a mean rate of 5.04 µg N g−1 day−1 during the predominantly wet condition in the forest. The overall nitrogen input from fixation was 6.1 kg N ha−1 year−1, of which 78% came from bryophytes, 18% from chlorolichens, and 4% from cyanolichens. Only 2.0% of the fixation occurred in cryptogams on the ground, whereas 67%, 24%, and 7% occurred on the lower branches, boles, and upper branches, respectively. These results show that tree epiphytes constitute a significant source of nitrogen for these forests, due to the trees’ large surface area, and can make up for the nitrogen lost from these ecosystems UCR::Vicerrectoría de Docencia::Ciencias Básicas::Facultad de Ciencias::Escuela de Biología UCR::Vicerrectoría de Investigación::Unidades de Investigación::Ciencias Básicas::Centro de Investigación en Biodiversidad y Ecología Tropical (CIBET)

Previous flux measurements in the perhumid cloud forest of northeastern Taiwan have shown efficient photosynthesis of the endemic tree species Chamaecyparis obtusa var. formosana even under foggy conditions in which leaf surface moisture would be expected. We hypothesized this to be the result of ‘xeromorphic’ traits of the Chamaecyparis leaves (hydrophobicity, stomatal crypts, stomatal clustering), which could prevent coverage of stomata by precipitation, fog, and condensation, thereby maintaining CO2 uptake. Here we studied the amount, distribution, and composition of moisture accumulated on Chamaecyparis leaf surfaces in situ in the cloud forest. We studied the effect of surface tension on gas penetration to stomata using optical O2 microelectrodes in the laboratory. We captured the dynamics of condensation to the leaf surfaces with an environmental scanning electron microscope (ESEM). In spite of substantial surface hydrophobicity, the mean water film thickness on branchlets under foggy conditions was 80 µm (upper surface) and 40 µm (lower surface). This amount of water could cover stomata and prevent CO2 uptake. This is avoided by the clustered arrangement of stomata within narrow clefts and the presence of Florin rings. These features keep stomatal pores free from water due to surface tension and provide efficient separation of plant and atmosphere in this perhumid environment. Air pollutants, particularly hygroscopic aerosol, may disturb this functionality by enhancing condensation and reducing the surface tension of leaf surface water.

Microclimate in the tropical montane cloud forest (TMCF) is variable on both spatial and temporal scales and can lead to large fluctuations in both leaf-level transpiration and whole plant water use. While variation in transpiration has been found in TMCFs, the influence of different microclimatic drivers on plant water relations in this ecosystem has been relatively understudied. Within the TMCF, epiphytes may be particularly affected by natural variation in microclimate due to their partial or complete disassociation from soil resources. In this study, we examined the effects of seasonal microclimate on whole plant water balance in epiphytes in both an observational and a manipulative experiment. We also evaluated the effects of different microclimatic drivers using three hierarchical linear (mixed) models. On average, 31 % of total positive sap flow was recovered via foliar water uptake (FWU) over the course of the study. We found that precipitation was the greatest driver of foliar water uptake and nighttime sap flow in our study species and that both VPD and precipitation were important drivers to daytime sap flow. We also found that despite adaptations to withstand seasonal drought, an extended dry period caused severe desiccation in most plants despite a large reduction in leaf-level and whole plant transpiration. Our results indicate that the epiphytes studied rely on FWU to maintain positive water balance in the dry season and that increases in dry periods in the TMCF may be detrimental to these common members of the epiphyte community.

Although the significance of canopy plant communities to ecosystem function is well documented, the process by which such communities become established in trees remains poorly known. Colonization of tree surfaces by canopy-dwelling plants often begins with the establishment of bryophytes, so the conditions that affect the dispersal of bryophytes in the forest canopy merit study. We assessed success rates of one mechanism of bryophyte propagation, the aerial dispersal of macroscopic fragments, using an experimental approach. We quantified interception and retention of marked fragments released from a 36 cm×36 cm grid 50 cm above branches of saplings and mature trees of the species Ocotea tonduzii in a montane cloud forest in Costa Rica. Only 1% of bryophyte fragments dropped over sapling crowns in this manner were retained for the 6-month duration of the study, while branches in the forest canopy with intact epiphyte loads and branches that had been stripped of their epiphytes retained 24% and 5%, respectively. Our results suggest that larger-diameter branches and the presence of other epiphytes can both improve the retention of bryophyte fragments on canopy branches. Further work will be needed to address the relative roles of other dispersal mechanisms (spores, gemmae, microscopic bryophyte fragments) and the dynamics of growth and establishment of macroscopic bryophyte fragments following their interception.

The physiological traits associated with water relations of eight common epiphytic ferns in a Mexican cloud forest were investigated in relation to the distribution of these species within the canopy. Fern distribution was significantly correlated with the relative water content at which stomata close, leaf thickness, stomatal density and size. Trichomanes bucinatum desiccated completely within hours in moderately dry air and was confined to the stem bases, and Asplenium cuspidatum, with no evident adaptations to cope with drought, grew in the second most shaded zone within the tree crowns. Despite growing in a humid cloud forest, all other species had xeric adaptations including coriaceous leaves (Pleopeltis mexicana, Elaphoglossum glaucum), succulent rhizomes (Polypodium puberulum, Phlebodium areolatum), low rates of uncontrolled water loss (all species except P. puberulum), leaf scales (Elaphoglossum petiolatum, Polypodium plebeium), and high cell wall elasticity (all species). P. plebeium and Pl. mexicanum, which grow in the most exposed locations, tolerated water loss beyond the turgor loss point before the stomata closed and appear to be poikilohydric or at least to tolerate high water deficits.

Climate warming predicts changes to the frequency and height of cloud-immersion events in mountain communities. Threatened southern Appalachian spruce-fir forests have been suggested to persist because of frequent periods of cloud immersion. These relic forests exist on only seven mountaintop areas, grow only above ca. 1,500 m elevation (maximum 2,037 m), and harbor the endemic Abies fraseri. To predict future distribution, we examined the ecophysiological effects of cloud immersion on saplings of A. fraseri and Picea rubens at their upper and lower elevational limits. Leaf photosynthesis, conductance, transpiration, xylem water potentials, and general abiotic variables were measured simultaneously on individuals at the top (1,960 m) and bottom (1,510 m) of their elevation limits on numerous clear and cloud-immersed days throughout the growing season. The high elevation sites had 1.5 as many cloud-immersed days (75 % of days) as the low elevation sites (56 % of days). Cloud immersion resulted in higher photosynthesis, leaf conductance, and xylem water potentials, particularly during afternoon measurements. Leaf conductance remained higher throughout the day with corresponding increases in photosynthesis and transpiration, despite low photon flux density levels, leading to an increase in water potentials from morning to afternoon. The endemic A. fraseri had a greater response in carbon gain and water balance in response to cloud immersion. Climate models predict warmer temperatures with a decrease in the frequency of cloud immersion for this region, leading to an environment on these peaks similar to elevations where spruce-fir communities currently do not exist. Because spruce-fir communities may rely on cloud immersion for improved carbon gain and water conservation, an upslope shift is likely if cloud ceilings rise. Their ultimate survival will likely depend on the magnitude of changes in cloud regimes.

A field manipulation experiment was conducted in a subtropical montane cloud forest in southwestern China to determine the possible responses of epiphytic bryophytes to increasing nitrogen (N) deposition from community to physiology level, and to find sensitive epiphytic bryophytes that may be used as indicators for assessing the degree of N pollution. N addition had significantly negative effects on species richness and cover of the epiphytic bryophyte community. Harmful effects of high N loads were recorded for chlorophyll, growth, and vitality of the species tested. The decline of some epiphytic bryophytes may result from detrimental effects on degradation to photosynthetic pigments. Bazzania himalayana (Mitt.) Schiffn., Bazzania ovistipula (Steph.) Mizut., and Homaliodendron flabellatum (Sm.) Fleisch. are candidates in atmospheric nitrogen monitoring. Epiphytic bryophytes in the montane cloud forest are very sensitive to increasing N deposition and often difficult to recover once they have been destroyed, providing early detection of enhanced N pollution for trees or even the whole forest ecosystem. The inference that increasing N pollution may lead to loss of biodiversity is a concern to the developing economy in western China, and should alert the government to the adverse impacts caused by increased industrial pollution during the process of China's West Development.

Few data are available describing the photosynthetic parameters of the leaves of tropical montane cloud forests (TMCF). Here, we present a study of photosynthetic leaf traits (V(cmax) and J(max)), foliar dark respiration (R(d)), foliar nitrogen (N) and phosphorus (P), and leaf mass per area (LMA) throughout the canopy for five different TMCF species at 3025 m a.s.l. in Andean Peru. All leaf traits showed a significant relationship with canopy height when expressed on an area basis, and V(cmax-area) and J(max-area) almost halved when descending through the TMCF canopy. When corrected to a common temperature, average V(cmax) and J(max) on a leaf area basis were similar to lowland tropical values, but lower when expressed on a mass basis, because of the higher TMCF LMA values. By contrast, R(d) on an area basis was higher than found in tropical lowland forests at a common temperature, and similar to lowland forests on a mass basis. The TMCF J(max)-V(cmax) relationship was steeper than in other tropical biomes, and we propose that this can be explained by either the light conditions or the relatively low VPD in the studied TMCF. Furthermore, V(cmax) had a significant-though relatively weak and shallow-relationship with N on an area basis, but not with P, which is consistent with the general hypothesis that TMCFs are N rather than P limited. Finally, the observed V(cmax)-N relationship (i.e., maximum photosynthetic nitrogen use efficiency) was distinctly different from those in tropical and temperate regions, probably because the TMCF leaves compensate for reduced Rubisco activity in cool environments.

Specific leaf area (SLA) is a key functional trait reflecting the trade-off between resource capture and conservation, and has been identified as playing an important role in plant community assembly. Mechanistic models of community assembly state that the assemblage of species in a local community is controlled by environment filters operating on functional traits. We measured within- and among-species variation of SLA, and environmental conditions in a tropical cloud forest to explore how variation in this functional trait contributes to community assembly. SLA variation at the species level was also decomposed into alpha (within assemblage variation), and beta (across assemblage variation) values. SLA decreased with increasing solar irradiance (approximated using plant height) within the three study sites, and differed among the three sites both for within- and among-species comparisons. Mean plot SLA, accounting for both within and among species across the three sites, increased significantly in relation to air temperature but not local photosynthetic photon flux density and soil total phosphorus. Alpha SLA decreased with increasing solar irradiance within the three sites and beta SLA differed among the three sites. Our results clearly demonstrate that light and air temperature are key environmental factors involved in organizing plant species within and among communities in tropical cloud forests. The strong relationship between both intra- and interspecific variation in SLA and environmental conditions strongly confirms the role of trait variation in the assembly of plant species in tropical cloud forest communities via environment filtering related to light availability and air temperature.

The red spruce-Fraser fir ecosystem [Picea rubens Sarg.-Abies fraseri (Pursh) Poir.] of the southern Appalachian mountains, USA, is a temperate zone cloud forest immersed in clouds for 30–40% of a typical summer day, and experiencing immersion on about 65% of all days annually. We compared the microclimate, photosynthetic gas exchange, and water relations of Fraser fir trees in open areas during cloud-immersed, low-cloud, or sunny periods. In contrast to sunny periods, cloud immersion reduced instantaneous sunlight irradiance by 10–50%, and midday atmospheric vapor pressure deficit (VPD) was 85% lower. Needle surfaces were wet for up to 16 h per day during cloud-immersed days compared to

The elevations at which tropical treelines occur are believed to represent the point where low mean temperatures limit the growth of upright woody trees. Consequently, tropical treelines are predicted to shift to higher elevations with global warming. However, treelines throughout the tropics have remained stationary despite increasing global mean temperatures. The goal of the study reported here was to build a more comprehensive understanding of the effects of mean temperature, low-temperature extremes, shading, and their interactions on seedling survival at tropical treelines. We conducted a seedling transplant study using three dominant canopy-forming treeline species in the southern tropical Andes. We found species-specific differences and contrasting responses in seedling survival to changes in mean temperature. The most abundant naturally occurring species at the seedling stage outside the treeline, Weinmannia fagaroides, showed a negative relationship between the survival of transplanted seedlings and mean temperature, the opposite of a priori expectations. Conversely, Clethra cuneata showed increased survival at higher mean temperatures, but survival also increased with higher absolute low temperatures and the presence of shade. Finally, the survival of Gynoxys nitida seedlings was insensitive to temperature but increased under shade. These findings show that multiple factors can determine the upper distributional limit of species forming the current tropical treeline. As such, predictions of future local and regional tropical treeline shifts may need to consider several factors beyond changes in mean temperature. If the treeline remains stationary and cloud forests are unable to expand into higher elevations, there may be severe species loss in this biodiversity hotspot.

The humid tropics represent only one example of the many places worldwide where anthropogenic disturbance and climate change are quickly affecting the feedbacks between water and trees. In this article, we address the need for a more long-term perspective on the effects of climate change on tropical montane cloud forests (TMCF) in order to fully assess the combined vulnerability and long-term response of tropical trees to changes in precipitation regimes, including cloud immersion. We first review the ecophysiological benefits that cloud water interception offers to trees in TMCF and then examine current climatological evidence that suggests changes in cloud base height and impending changes in cloud immersion for TMCF. Finally, we propose an experimental approach to examine the long-term dynamics of tropical trees in TMCF in response to environmental conditions on decade-to-century time scales. This information is important to assess the vulnerability and long-term response of TMCF to changes in cloud cover and fog frequency and duration.

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.

Fog has been viewed as an important source of moisture in many coastal ecosystems, yet its impor- tance 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 oA the redwood trees themselves (interception input). When trees were absent, the average annual input from fog was only 17%, demonstrating that the trees significantly in- fluence the magnitude of fog water input to the ecosys- tem. Stable hydrogen and oxygen isotope analyses of water from fog, rain, soil water, and xylem water ex- tracted 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 fre- quent, 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, es- pecially in summer when rainfall was absent. Depen- dence on fog as a moisture source was highest in the year when rainfall was lowest but fog inputs normal. Inter- estingly, during the mild El Nino year of 1993, when the ratio of rainfall to fog water input was significantly higher and fog inputs were lower, both the proportion and coeAcient 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 Nino 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 eco- systems and the plants which inhabit them.

The relative advantages of being deciduous or evergreen in subtropical forests and the relationship between leaf phenology and nutrient resorption efficiency are not well understood. The most successful deciduous species (Lyonia ovalifolia) in an evergreen-dominated subtropical montane cloud forest in southwest (SW) China maintains red senescing leaves throughout much of the winter. The aim of this study was to investigate whether red senescing leaves of this species were able to assimilate carbon in winter, to infer the importance of maintaining a positive winter carbon balance in subtropical forests, and to test whether an extended leaf life span is associated with enhanced nutrient resorption and yearly carbon gain. The red senescing leaves of L. ovalifolia assimilated considerable carbon during part of the winter, resulting in a higher yearly carbon gain than cooccurring deciduous species. Its leaf N and P resorption efficiency was higher than for co-occurring non-anthocyanic deciduous species that dropped leaves in autumn, supporting the hypothesis that anthocyanin accumulation and/or extended leaf senescence help in nutrient resorption. Substantial winter carbon gain and efficient nutrient resorption may partially explain the success of L. ovalifolia versus that of the other deciduous species in this subtropical forest. The importance of maintaining a positive carbon balance for ecological success in this forest also provides indirect evidence for the dominance of evergreen species in the subtropical forests of SW China. Fil: Zhang, Yong Jiang. Chinese Academy Of Sciences; República de China Fil: Yang, Qiu Yun. Chinese Academy Of Sciences; República de China Fil: Lee, David W.. Florida International University; Estados Unidos Fil: Goldstein, Guillermo Hernan. University of Miami; Estados Unidos. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Cao, Kun Fang. Chinese Academy Of Sciences; República de China

From studies in seasonal lowland tropical forests, bromeliad epiphytes appear to be limited mainly by water, and to a lesser extent by nutrient supply, especially phosphorous. Less is understood about the mineral nutrition of tropical montane cloud forest (TMCF) epiphytes, even though their highest diversity is in this habitat. Nutrient limitation is known to be a key factor restricting forest productivity in TMCF, and if epiphytes are nutritionally linked to their host trees, as has been suggested, we would expect that they are also nutrient limited. We studied the effect of a higher nutrient input on reproduction and growth of the tank bromeliad Werauhia sintenisii in experimental plots located in a TMCF in Puerto Rico, where all macro- and micronutrients had been added quarterly starting in 1989 and continuing throughout the duration of this study. We found that bromeliads growing in fertilized plots were receiving litterfall with higher concentrations of N, P, and Zn and had higher concentrations of P, Zn, Fe, Al, and Na in their vegetative body. The N:P ratios found (fertilized = 27.5 and non-fertilized = 33.8) suggest that W. sintenisii may also be phosphorous limited as are lowland epiphytes. Fertilized plants had slightly longer inflorescences, and more flowers per inflorescence, than non-fertilized plants, but their flowers produced nectar in similar concentrations and quantities. Fertilized plants produced more seeds per fruit and per plant. Frequency of flowering in two consecutive years was higher for fertilized plants than for controls, suggesting that fertilized plants overcome the cost of reproduction more readily than non-fertilized plants. These results provide evidence that TMCF epiphytic bromeliads are nutrient limited like their lowland counterparts.

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.

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.