Responses of a common tropical epiphyte, Asplenium nidus, to changes in water and nutrient availability.

Epiphytes are highly dependent on atmospheric inputs of water and nutrients. Reductions in water availability associated with warming and climate change and continual atmospheric nitrogen (N) deposition can affect plant growth but few studies have evaluated the effects of changes in both water and nutrient availabilities on epiphytes. We experimentally tested whether epiphyte growth is more water- or nutrient-limited, if nutrient limitation was stronger for nitrogen or phosphorus, and whether nutrient limitation interacts with water availability. We applied watering (high and low) and nutrient addition (control, +N, +P, +N+P) treatments to greenhouse-grown Asplenium nidus, a common epiphytic fern found in many tropical and subtropical wet forests. We measured leaf area production and leaf elemental concentrations to assess how A. nidus growth and physiology respond to changes in water and nutrient availabilities. We found that leaf growth of A. nidus was more affected by water availability than nutrient addition and the effect of adding nutrients was not fully realized under low-water availability. Among the different nutrient treatments, +N+P had the greatest effects on A. nidus growth and physiology in both watering treatments. Watering treatment changed leaf elemental concentrations but not their ratios (i.e. C:N and N:P). Nutrient addition altered C:N and N:P ratios and increased the concentration of the added elements in leaves, with more pronounced increases in the high-watering treatment. We conclude that the growth of A. nidus is more water- than nutrient-limited. When nutrient limitation occurs (i.e. under high-water availability), nutrient co-limitation is stronger than limitation by N or P alone. This result taken together with studies of other epiphytes suggests greater water than nutrient limitation is likely widespread among epiphytic plants. The limited effects of nutrient addition in the low-water treatment suggest that the effect of atmospheric N deposition on epiphyte growth will be limited when water availability is low.

Fog and rain water chemistry in a tea plantation of northern Taiwan.

Tea plantations are expanding globally and many are in mountainous areas with frequent fog but few studies have examined fog chemistry in these areas. We examined chemical composition of fog and rain water at a tea plantation in northern Taiwan. Fog water was collected using a Kroneis passive cylindrical fog-water collector and rain water was collected using a 20-cm-diameter funnel. The most abundant ions were Cl- and Na+ in both fog and rain waters due to the proximity of the site to the coast. The order of abundance of other ions was NO3- > Mg2+ > SO42- > Ca2+ > NH4+ > K+ > H+ in fog water and SO42- > K+ > NO3- > NH4+ > Ca2+ > Mg2+ > H+ in rain water. The concentration enrichment ratio (fog to rain) ranged between 2.2 (K+) and 22 (Mg2+) lying between sites near major emission sources and sites in remote areas, possibly because the immediate surrounding landscape is covered with secondary forests although it is near large cities. Factor analysis highlights the influences of sea-salt aerosols on the variation of fog and rain water chemistry. Sea-salt corrections using Na+ as the sea salt tracer led to negative concentrations of Cl- and Mg2+ suggesting that assumptions involved in sea-salt corrections were not satisfied. Agriculture influence is identified as a unique factor for explaining variance of K+, NH4+, and dissolved organic nitrogen (DON) concentrations in fog water but not rain water. Ion concentrations in fog and rain water were generally higher in the weekly samples associated with air trajectories passing through the continental East Asia than those associated with oceanic trajectories pointing to the role of regional pollution sources in affecting local fog and rain water chemistry. Our study highlights greater effects of tea agriculture on fog than rain water chemistry.

Air pollutant removal by four sidewalk tree species in the largest city in Taiwan.

Air pollutants pose risks to human health, especially in densely populated cities. We compared the interception of suspended particles and metal elements by four sidewalk tree species with different leaf surface wettability (based on contact angle), leaf area, and phenology in Taipei, Taiwan. Suspended particles were enriched 2.0-2.5 times in throughfall relative to rainfall due to wash-off of suspended particles deposited on leaf surfaces during rainless periods. The enrichment in throughfall was greater in tree species with larger leaf areas. Despite greater concentrations of suspended particles in rainfall during the low-leaf-area period, enrichment was greater in the high-leaf-area period, indicating that leaf area was a key factor affecting canopy interception of pollutants. Throughfall enrichment of suspended particles positively correlated with water quantity, indicating that air pollutants intercepted by tree canopies were not fully washed off by rainfall. Annually, ∼830 g of suspended particles were intercepted and washed off from one tree canopy, with a crown area of 42 m2 . Scaling up, a rough estimate of 72.7 Mg of suspended particles were intercepted annually by the 90,000 sidewalk trees in Taipei City. Copper, chromium, and aluminum were enriched in throughfall compared with rainfall. However, lead was depleted in throughfall, indicating greater interception than wash-off. Based on our results, leaf area and length of foliated period are key characteristics affecting canopy interception of particulate matter and associated metal elements, whereas leaf surface wettability is of secondary importance.

A general pattern of trade-offs between ecosystem resistance and resilience to tropical cyclones.

Tropical cyclones drive coastal ecosystem dynamics, and their frequency, intensity, and spatial distribution are predicted to shift with climate change. Patterns of resistance and resilience were synthesized for 4138 ecosystem time series from n = 26 storms occurring between 1985 and 2018 in the Northern Hemisphere to predict how coastal ecosystems will respond to future disturbance regimes. Data were grouped by ecosystems (fresh water, salt water, terrestrial, and wetland) and response categories (biogeochemistry, hydrography, mobile biota, sedentary fauna, and vascular plants). We observed a repeated pattern of trade-offs between resistance and resilience across analyses. These patterns are likely the outcomes of evolutionary adaptation, they conform to disturbance theories, and they indicate that consistent rules may govern ecosystem susceptibility to tropical cyclones.

Changes of precipitation acidity related to sulfur and nitrogen deposition in forests across three continents in north hemisphere over last two decades.

Through synthesizing bulk precipitation chemistry in forest sites (n = 128) from three monitoring networks, (NADP in Northern America, EMEP in Europe, and EANET in East Asia), this study quantifies the temporal changes of precipitation acidity and its dominant acidifying agents over the last two decades. Results show distinct declines of sulfate and nitrate depositions and increases of precipitation pH in northeast America and central and east Europe, but not in Asia during 1999 and 2018. The decreases of sulfate and nitrate depositions likely reflect the long-term effort of pollutant emission controls. The temporal pattern of sulfate (SO42-)/nitrate (NO3-) and ammonium nitrogen (NH4-N)/nitrate nitrogen (NO3-N) equivalent ratios indicate that acid rain in the NADP and EMEP have transitioned from sulfate-dominated to nitrate-dominated, and the DIN deposition has shifted from nitrate-dominated to ammonium-dominated in recent years, owing to reductions of sulfur dioxides (SO2) and nitrogen oxides (NOx) emissions. In contrast, sulfate still plays a dominant role on the acidity of precipitation than nitrate in Asia, and NH4-N deposition also has a significant contribution in N flux due to increasing trends of ammonia emissions in Southeast Asia.

Tropical Cyclone Ecology: A Scale-Link Perspective.

Tropical cyclones are increasing in intensity and size and, thus, are poised to increase in importance as disturbance agents. Our understanding of cyclone ecology is biased towards the North Atlantic Basin, because cyclone effects do differ across oceanic basins. Cyclones have both short and long-term effects across the levels of biological organization, but we lack a scale-perspective of cyclone ecology. Effects on individual trees, such as defoliation or branch stripping and uprooting, are mechanistically linked to effects at the community and ecosystem levels, including forest productivity and stand regeneration time. Forest dwarfing via the gradual removal of taller trees by cyclones over many generations illustrates that cyclones shape forest structure through the accumulation of short-term effects over longer timescales.

Temporal changes in soil C-N-P stoichiometry over the past 60 years across subtropical China.

Controlled experiments have shown that global changes decouple the biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P), resulting in shifting stoichiometry that lies at the core of ecosystem functioning. However, the response of soil stoichiometry to global changes in natural ecosystems with different soil depths, vegetation types, and climate gradients remains poorly understood. Based on 2,736 observations along soil profiles of 0-150 cm depth from 1955 to 2016, we evaluated the temporal changes in soil C-N-P stoichiometry across subtropical China, where soils are P-impoverished, with diverse vegetation, soil, and parent material types and a wide range of climate gradients. We found a significant overall increase in soil total C concentration and a decrease in soil total P concentration, resulting in increasing soil C:P and N:P ratios during the past 60 years across all soil depths. Although average soil N concentration did not change, soil C:N increased in topsoil while decreasing in deeper soil. The temporal trends in soil C-N-P stoichiometry differed among vegetation, soil, parent material types, and spatial climate variations, with significantly increased C:P and N:P ratios for evergreen broadleaf forest and highly weathered Ultisols, and more pronounced temporal changes in soil C:N, N:P, and C:P ratios at low elevations. Our sensitivity analysis suggests that the temporal changes in soil stoichiometry resulted from elevated N deposition, rising atmospheric CO2 concentration and regional warming. Our findings revealed that the responses of soil C-N-P and stoichiometry to long-term global changes have occurred across the whole soil depth in subtropical China and the magnitudes of the changes in soil stoichiometry are dependent on vegetation types, soil types, and spatial climate variations.

Impacts of increasing typhoons on the structure and function of a subtropical forest: reflections of a changing climate.

Due to their destructive and sporadic nature, it is often difficult to evaluate and predict the effects of typhoon on forest ecosystem patterns and processes. We used a 21-yr record of litterfall rates to explore the influence of typhoon frequency and intensity, along with other meteorological variables, on ecosystem dynamics in a subtropical rainforest. Over the past half century there has been an increasing frequency of strong typhoons (category 3; >49.6 m s-1; increase of 1.5 typhoons/decade) impacting the Fushan Experimental Forest, Taiwan. At Fushan strong typhoons drive total litterfall mass with an average of 1100 kg ha-1 litterfall typhoon-1. While mean typhoon season litterfall has been observed to vary by an order of magnitude, mean litterfall rates associated with annual leaf senescence vary by <20%. In response to increasing typhoon frequency, total annual litter mass increased gradually over the 21-year record following three major typhoons in 1994. Monthly maximum wind speed was predictive of monthly litterfall, yet the influence of precipitation and temperature was only evident in non-typhoon affected months. The response of this subtropical forest to strong typhoons suggests that increasing typhoon frequency has already shifted ecosystem structure and function (declining carbon sequestration and forest stature).

Trends of two decadal precipitation chemistry in a subtropical rainforest in East Asia.

Long-term monitoring of precipitation chemistry provides a great opportunity to examine the evolution of air pollutant emissions and effectiveness of air pollution control measures. We evaluated the characteristics and trends of precipitation chemistry at both annual and seasonal scales based on the records of 1994-2013 at Fushan Experimental Forest (FEF) of northeastern Taiwan. The results showed that 77% of the weekly precipitation had pH<5.0. The two-decadal average annual pH was 4.62, without a significant inter-annual trend, possibly due to the concurrent declines of both acidic pollutants and base cations. There was a significant positive relationship between [SO42-+NO3-] and [Ca2++NH4+] indicating that their deposition was likely dominated by NH4NO3, (NH4)2SO4, Ca(NO3)2, and CaSO4. There was a significant negative relationship between precipitation pH and the difference between [SO42-+NO3-] and [Ca2++NH4+], not just [SO42-+NO3-], suggesting that precipitation acidity was not solely determined by acidic pollutants but by the balance between acidic pollutants and base cations. We also found temporal decreases of Ca2+ and NH4+ concentrations in precipitation which contributed to the low acid neutralization capacity of precipitation. Annual deposition of NO3- and SO42- was 23 and 55kgha-1yr-1, which is much higher than most forest sites in the industrialized countries suggesting that acid deposition is still a major environmental issue in Taiwan. Annual deposition of NH4+, Ca2+ and NO3- showed significant decreasing trends during the 20-year period, which was mostly due to the decreases in the summer deposition associated with air pollution mitigation strategies. Winter deposition showed no decreasing patterns for the same period. The high contribution to annual acid deposition from autumn-winter and spring rains (50%) associated with northeast monsoon implies that long-range transport of anthropogenic emissions from East Asia played a key role on acid depositions at FEF and possibly many areas in the region. Therefore, intergovernmental cooperation is urgently needed to effectively mitigate the threat of acid deposition in East Asia.

Modeling the terrestrial N processes in a small mountain catchment through INCA-N: A case study in Taiwan.

Riverine dissolved inorganic nitrogen (DIN) is an important indicator of trophic status of aquatic ecosystems. High riverine DIN export in Taiwan, ~3800kg-Nkm-2yr-1, which is ~18 times higher than the global average, urges the need of thorough understanding of N cycling processes. We applied INCA-N (Integrated Nitrogen Catchment Model) to simulate riverine DIN export and infer terrestrial N processes using weekly rainwater and streamwater samples collected at the Fushan Experimental Forest (FEF) of northern Taiwan. Results showed that the modeled discharge and nitrate export are in good agreement with observations, suggesting the validity of our application. Based on our modeling, the three main N removal processes, in the order of descending importance, were plant uptake, riverine N transport and denitrification at FEF. The high plant uptake rate, 4920kg-Nkm-2yr-1, should have led to accumulation of large biomass but biomass at FEF was relatively small compared to other tropical forests, likely due to periodic typhoon disruptions. The low nitrate concentration but high DIN export highlights the importance of hydrological control over DIN export, particularly during typhoons. The denitrification rate, 750kg-Nkm-2yr-1, at FEF was also low compared to other tropical forest ecosystems, likely resulting from quick water drainage through the coarse-loamy top soils. The high DIN export to atmospheric deposition ratio, 0.45, suggests that FEF may be in advanced stages of N excess. This simulation provides useful insights for establishing monitoring programs and improves our understanding N cycling in subtropical watersheds.

Effects of different N sources on riverine DIN export and retention in a subtropical high-standing island, Taiwan.

Increases in nitrogen (N) availability and mobility resulting from anthropogenic activities have substantially altered the N cycle, both locally and globally. Taiwan characterized by the subtropical montane landscape with abundant rainfall, downwind of the most rapidly industrializing eastern coast of China, can be a demonstration site for extremely high N input and riverine DIN (dissolved inorganic N) export. We used 49 watersheds with similar climatic and landscape settings but classified into low, moderate, and highly disturbed categories based on population density to illustrate their differences in nitrogen inputs (through atmospheric N deposition, synthetic fertilizers, and human emission) and DIN export ratios. Our results showed that the island-wide average riverine DIN export is ~ 3800 kg N km-2 yr-1, approximately 18 times the global average. The average riverine DIN export ratios are 0.30-0.51, which are much higher than the averages of 0.20-0.25 of large rivers around the world, indicating excessive N input relative to ecosystem demand or retention capacity. The low disturbed watersheds have a high N retention capacity and DIN export ratios of 0.06-0.18 in spite of the high N input (~ 4900 kg N km-2 yr-1). The high retention capacity is likely due to effective uptake by secondary forests in the watersheds. The moderately disturbed watersheds show a linear increase in DIN export with increases in total N inputs and mean DIN export ratios of 0.20 to 0.31. The main difference in land use between low and moderately disturbed watersheds is the greater proportion of agricultural land cover in the moderately disturbed watersheds. Thus, their greater DIN export could be attributed to N fertilizers used in the agricultural lands. The greater export ratios also imply that agricultural lands have a lower proportional N retention capacity and that reforestation could be an effective land management practice to reduce riverine DIN export. The export ratios of the highly disturbed watersheds are very high, 0.42-0.53, suggesting that much of the N input is transported downstream directly, and urges the need to increase the proportion of households connected to a sewage system and improve the effectiveness of wastewater treatment systems. The increases in the riverine DIN export ratio along the gradient of human disturbance also suggest a gradient in N saturation in subtropical Taiwan. Our results help to improve our understanding of factors controlling riverine DIN export and provide empirical evidence that calls for sound N emission/pollution control measures.

Ecological facilitation between two epiphytes through drought mitigation in a subtropical rainforest.

Positive species interactions (facilitation) play an important role in shaping the structures and species diversity of ecological communities, particularly under stressful environmental conditions. Epiphytes in rainforests often grow in multiple-species clumps, suggesting interspecies facilitation. However, little is known about the patterns and mechanisms of epiphyte co-occurrence. We assessed the interactions of two widespread epiphyte species, Asplenium antiquum and Haplopteris zosterifolia, by examining their co-occurrence and size-class association in the field. To elucidate factors controlling their interactions, we conducted reciprocal-removal and greenhouse-drought experiments, and nutrient and isotope analyses. Forty-five percent of H. zosterifolia co-occurred with A. antiquum, whereas only 17% of A. antiquum co-occurred with H. zosterifolia. Removing the fronds plus substrate of A. antiquum reduced the relative frond length and specific leaf area of H. zosterifolia, but removing fronds only had little effect. Removing H. zosterifolia had no significant effects on the growth of A. antiquum. H. zosterifolia co-occurring and not co-occurring with A. antiquum had similar foliar nutrient concentrations and δ(15)N values, suggesting that A. antiquum does not affect the nutrient status of H. zosterifolia. Reduced growth of H. zosterifolia with the removal of A. antiquum substrate, together with higher foliar δ(13)C for H. zosterifolia growing alone than those co-occurring with A. antiquum, suggest that A. antiquum enhances water availability to H. zosterifolia. This enhancement probably resulted from water storage in the substrate of A. antiquum, which could hold water up to 6.2 times its dry weight, and from reduced evapotranspiration due to shading of A. antiquum fronds. Greater water loss occurred in the frond-clipped group than the unclipped group between days 3-13 of the drought treatment. Our results imply that drought mitigation by substrate-forming epiphytes is important for maintaining epiphyte diversity in tropic and subtropic regions with episodic water limitations, especially in the context of anthropogenic climate change.

Fog and precipitation chemistry at a mid-land forest in central Taiwan.

We analyzed fog and bulk precipitation chemistry at a cloud forest in central Taiwan where mountain agriculture activities are highest. There were 320 foggy days (visibility <1000 m) recorded between April 2005 and March 2006. Fog was most frequent between April 2005 and July 2005 and in March 2006 (153/153 d) and least frequent in January 2006 (21/31 d). The total fog duration was 2415 h, representing 28% of the sampling period. Compared with bulk precipitation, fog was disproportionally enriched in NO(3)(-) and SO(4)(2-) relative to K(+), Ca(2+), Mg(2+), and NH(4)(+), resulting in higher a content of nitric acid and sulfuric acid than weak acids or neutral salts and, therefore, higher acidity (median pH, 4.9) in fog than in bulk precipitation (median and mean pH, 5.5). The very high input of NH(4)(+) (47 kg N ha(-1) yr(-1)) through bulk precipitation suggests that the use of fertilizer (ammonium sulfate and animal manure) associated with mountain agriculture has a major impact on atmospheric deposition at the surrounding forest ecosystems. The input of inorganic N reached 125 kg N ha(-1) yr(-1) and likely exceeded the biological demand of the forest ecosystem. Sulfate is the most abundant anion in fog at Chi-tou and in precipitation at various forests throughout Taiwan, suggesting that the emission and transport of large quantities of SO(2,) the precursor of SO(4)(2-), is an island-wide environmental issue.