Greenfall leaves caused by hailstorms may represent a resource pulse of nutrients. We determined the contribution of greenfall versus senescent leaves to total litterfall production, carbon, nitrogen and phosphorus input to the system, and leaf decomposition rate. Litterfall was collected monthly for three years in two cloud forests (F1, F2) and a restoration planting area (R) in Veracruz, Mexico. Two fortuitous hailstorms occurred in the second year. Leaf decomposition rate was determined in all three sites but did not differ across them. Total annual litterfall, excluding greenfall, was 10.0, 10.1, and 7.7 Mg ha−1 y−1 for F1, F2, and R, respectively. Senescent leaves represented 65% of the litterfall, while greenfall leaves increased the annual leaf biomass component of the litterfall by 12%. Concentrations of carbon, nitrogen, and phosphorus were 2.3, 5.7, and 18.1% higher, respectively, in greenfall than in senescent leaves. Greenfall increased the annual input of C, N, and P by 12, 13, and 14%, respectively. Despite their short duration (approximately 70 minutes), the hailstorm events generated a substantial contribution of greenfall leaves and a source of extra C, N, and P, since these leaves decompose and are incorporated into the cloud forest system.

Litterfall is one of the most important pathways through which nutrients are recycled in the terrestrial biosphere. In tropical soils, which are generally low in essential nutrients such as phosphorus and cations, the flux of nutrients through litterfall is particularly important to sustaining CO2-uptake capacity; however, questions remain over the role of altitude in altering litter nutrient cycling rates among tropical forest ecosystems. Here we examine litterfall, carbon (C), nitrogen (N) and phosphorus (P) fluxes through litterfall over an altitudinal gradient in the coastal Atlantic Forest located on the northern coast of the State of São Paulo, Brazil. Litterfall was collected twice a month for 1 y (April 2007–March 2008) using 30 litter traps placed in four different forest types arrayed by altitude – coastal forest (sea level), lowland forest (50–200 m asl), submontane forest (300–500 m asl) and montane forest (1000 m asl). Litterfall mass-fluxes decreased with increasing altitude, from ~9 Mg ha−1 in lowland forests to 7 Mg ha−1 in higher-altitude ecosystems. Contribution of reproductive organs to litterfall was significantly greater in lower than in higher altitudes. Litterfall N and P fluxes were higher in the lowland forest vs. other forest types, pointing to strong altitudinal controls over nutrient cycling. Furthermore, nitrogen-use efficiency (NUE) was lower and litter δ15N was higher in the lowland site providing additional evidence for lack of N constraints to productivity in lowland of the south-eastern Atlantic Forest.

Solar irradiance and precipitation are the most likely drivers of the seasonal variation of net primary productivity (NPP) in tropical forests. Since their roles remain poorly understood, we use litter traps, dendrometer bands and census data collected from one hectare permanent plots to quantify the seasonality of above-ground NPP components and weather parameters in 13 sites distributed along a 2800-m altitudinal gradient ranging from lowland Amazonia to the high Andes. We combine canopy leaf area index and litterfall data to describe the seasonality of canopy production. We hypothesize that solar irradiance is the primary driver of canopy phenology in wetter sites, whereas precipitation drives phenology in drier systems. The seasonal rhythm of canopy NPP components is in synchrony with solar irradiance at all altitudes. Leaf litterfall peaks in the late dry season, both in lowland (averaging 0.54 ± 0.08 Mg C ha y−1, n = 5) and montane forests (averaging 0.29 ± 0.04 Mg C ha y−1, n = 8). Peaks in above-ground coarse woody NPP appears to be triggered by the onset of rainfall in seasonal lowland rain forests (averaging 0.26 ± 0.04 Mg C ha y−1, n = 5, in November), but not in montane cloud forests.

High densities of palms are common in many tropical forests. In some cases, the dominance of palms has been associated with a depauperate understorey and high rates of native seedling mortality. A variety of different potential mechanisms has been suggested to explain the sustained palm dominance in the understorey and canopy of these forests. Working in a Cocos nucifera-dominated wet tropical forest at Palmyra Atoll in the central Pacific, we examine how litterfall from this pantropical, and economically important palm, impacts seedling survival. We compare rates of litterfall, and rates of litterfall-associated damage, between forest stands dominated by C. nucifera (coconut palm) and forest stands with low abundance of C. nucifera. To assess litterfall damage we survey damage to both artificial seedlings (n = 711), outplanted real seedlings of two species (with and without protection via caging; n = 204), and standing rates of litterfall damage. We find that rates of large-litterfall damage were an average of five times higher in sites with high densities of C. nucifera. Associated with these increases we observe that levels of physical damage to artificial model seedlings caused by litterfall over a 4-mo period increased from 4.9% in sites with low abundance of C. nucifera to 16.1% in sites with high abundance of C. nucifera. Extrapolated to annual rates, litterfall damage of this magnitude exceeds the average levels observed in other published studies. Living native seedlings also showed more than 300% higher levels of mortality in forest stands with high densities of C. nucifera, a difference that was greatly reduced when protected by caging from litterfall. In contrast, uncaged C. nucifera seedlings actually had slightly higher survivorship in habitats dominated by conspecifics. We suggest that litterfall damage may be an important mechanism by which this tropical palm reaches and maintains near monodominance in many coastal and insular habitats.

In heterotrophic streams the retention and export of coarse particulate organic matter and associated elements are fundamental biogeochemical processes that influence water quality, food webs and the structural complexity of forested headwater streams. Nevertheless, few studies have documented the quantity and quality of exported organic matter over multiple years and under a range of conditions that includes both droughts and hurricanes. This study quantifies the export of coarse particulate organic matter (CPOM, > 12.7 mm), over 18 y in two headwater streams in north-east Puerto Rico. Daily exports ranged from 0 to over 170 g ha−1 d−1 and averaged 7.39 g ha−1 d−1, with similar amounts coming from leaves (3.5 g ha−1 d−1) and wood (3.2 g ha−1 d−1). Export of coarse particulate organic carbon was 3.0 g ha−1 d−1 which constitutes only 1.32% of carbon exports. Most litter falling into the streams was processed in place as only 2.3% of the leaf litter falling directly into these perennial channels was exported as CPOM. On average, 6 wk y−1 had no exports while events transporting more than 10 g ha−1 d−1 occurred every 2.8 mo. Instead of a single annual pulse as observed in deciduous systems, there were annual peaks in CPOM exports during May and September and less export during the drier period from December to February. Ratios of C:N in the exported material were highest in the driest month and lowest during rainy months, while leaf fluxes for nitrogen, phosphorus and calcium were highest in rainy months and lowest during February. Although median daily exports and exports during low- and base-flow periods were similar before and after Hugo, after 16 y exports during moderate- and high-flow periods were still less than those in the 2 y prior to the hurricane. Our observations indicate a system with high rates of internal processing that quickly returns to median daily conditions following hurricanes but requires several decades for storm-flow exports to return to pre-disturbance conditions and indicates that the long-term pattern of CPOM export is associated with the level of maturity of watershed vegetation.

To evaluate N limitation on above-ground net primary production in a tropical lower montane rain forest, an N fertilization experiment was conducted for 2 y. The study site is located at 1200–1300 m asl in the Fortuna forest reserve in western Panama and has a mature, mixed-species stand growing on an Andisol soil. Control and N-fertilized (125 kg urea-N ha−1 y−1) treatments were represented by four replicate plots (each 40 × 40 m, separated by at least 40 m). Stem diameter growth was analysed by diameter at breast height classes and also for the three most abundant species. The three species did not respond to N addition. The response of stem growth and above-ground woody biomass production to N fertilization varied among dbh classes. Stem growth of trees of 10–30 cm dbh increased only in the first year of N addition while trees of 30–50 cm dbh responded in the second year of N addition, which may be due to differences in light conditions between years. Trees >50 cm dbh did not respond during 2 years of N addition. As a result, the overall stem growth and above-ground woody biomass production were not affected by N fertilization. Annual total fine litterfall increased in the first year of N fertilization, while annual leaf litterfall increased in both years of N addition. Above-ground net primary production, of which total fine litterfall constituted 68%, also increased only in the first year of N addition. The magnitude and timing of response of stem diameter growth and litterfall suggest that these aspects of above-ground productivity are not uniformly limited by N availability.

Analysis of litterfall, nutrient return and floor mass dynamics was made in mature and recently harvested bamboo savanna sites in a dry tropical region in India. The total annual litterfall was 590 and 267 g m–2 in mature and harvested sites, respectively. Of the total litterfall, 58–77% by weight was contributed by leaf litter. Although litterfall occurred throughout the year, as much as 72–83% of the leaf litterfall was recorded during winter (December–February). Seasonal distribution of total litterfall was: rainy 12–22% of annual, winter 64–65% and summer 13–22%. Nutrient concentrations in bamboo leaf litter were low compared with the other species growing in the same region. Annual nutrient return through litterfall amounted to: 28–49 kg N ha–1, 1.32.9 kg P ha–1, 33–85 kg Ca ha–1, 12–32 kg K ha–1 and 1.4–2.8 kg Na ha–1. While maximum accumulations of fresh litter on the ground occurred in summer, the partly decayed and particulate litter categories peaked in the rainy season. Decaying litter categories showed 1.5–3 times greater nutrient concentrations than the concentrations in fresh leaf litter. Marked immobilization of N and P occurred in decomposing leaf and particulate litter. The mean annual nutrient content of the floor was (kg ha–1): N 29–40, P 1.0–1.9, Ca 13.8–23.9, K 14.6–15.3 and Na 0.85–0.99. The annual turnover in litter layer was: 63–81% for dry mass and 64–90% for different nutrients.

Damage to Hawaiian Acacia koa forest by Hurricane Iniki was assessed by comparison with our previous measures of stand structure and leaf area index (LAI) at sites along a precipitation/elevation gradient on western Kauai. Reductions in LAI ranged from 29 to 80% and were correlated with pre-hurricane LAI and canopy height. The canopy damage resulted in a large pulse of litter, ranging from 4 to 19 t ha-1 across our study sites. In the first year post-hurricane, LAI recovery and relative increase in stand basal area were negatively correlated with the fraction of canopy LAI lost. The two major overstorey species, A. koa and Metrosiderous polymorpha, were damaged less than the sub-canopy species, Dodonaea viscosa and Psidium guajava. Although D. viscosa and P. guajava were both severely damaged in the hurricane, the alien species P. guajava had much higher survival than the native D. viscosa, both as adults and as seedlings. However, seedling density of P. guajava was limited by low recruitment. At these sites, there was no drastic change in species composition following hurricane disturbance, and forest structure and productivity had recovered to a great degree within 2 y.

Phenological observations were made in a Sumatran rain forest during three years (1980–1982). Phenological changes followed a consistent seasonal pattern. The abundance of young leaves and the fall of leaf litter peaked between December and February (first dry season); flowers were most abundant between January and April (first dry and first wet sea son), and ripe fruits in July-August (the second dry season). The fruit of strangling fig trees showed peaks in April and October, both wet season months. Within the study area there was variation in both the phase and the amplitude of the phenological cycles. One year, 1981, displayed mast flowering and fruiting. The observations indicate that the conditions for production were better during the mast year, a finding that facilitates our understanding of the evolution of mast fruiting.

Litterfall was measured monthly in ten 1 m2 traps in each of two 0.25 ha plots in the rain forest of Southern Bakundu Forest Reserve, Cameroon for 18 months from 1 July 1982 to 31 December 1983. Particular emphasis was placed on measuring species contributions and in relating litterfall to climatic factors; the litterfall data were also used in estimating leaf production. Annual litterfall ranged between 12.9 and 14.1 t ha−1 and was partitioned to leaves (61–66%), branches (23–38%), fruits and flowers (8–12%) and ephiphytes (0.8%). Emergents and top canopy species like Desbordesia glaucescens (evergreen) and Ceiba pentandra (deciduous) contributed more litter than the numerically dominant lower storey species like Cola lepidota and Diospyros spp. Litter fell throughout the year but was greatest during the dry season (November to March); litterfall was thus strongly negatively correlated with environmental moisture variables and could be predicted from its linear regression on time of year, throughfall and relative humidity. By correcting the leaf fall data for leaf weight losses due to grazing and re-translocation, leaf production rates of 10.2 to 11.2 t ha−1 yr−1 were estimated for the forest.

An investigation of seasonal rates of litter production in a mixed forest, Hong Kong Island, was undertaken from October 1982 to January 1984. Total annual litterfall amounted to 1218.96 g m−2 yr−1, comprising 68.6% leaves, 18.9% woody material, and 12.5% fruits, seeds and insect frass. 398.70 g m−2 of litter was recorded immediately after a severe typhoon (9 September 1983). Rates of leaf-fall and organic debris production were correlated with prevailing temperature and rainfall. Leaf-fall peaked in January 1983, March to July 1983, and in September 1983 after the typhoon, while large amounts of insect faeces were collected in May and June. Woody litter production rates were correlated with temperature and rainfall recorded in previous months. Woody litterfall exhibited no obvious periodicity but was slightly increased during March and April. The seasonal distribution of litterfall is discussed in relation to climatic factors and habitat nutrient economy.

Rates of litterfall over a period of 3 y at one undisturbed site and two sites disturbed by selective harvesting are reported for a tropical rain forest area in North Queensland, Australia. Litterfall over a period of 18 months is also reported for a further undisturbed site in the same area. No significant differences were found in annual litterfall between the sites, with annual litterfall rates ranging from 5.0 to 6.0 t ha−1 y−1. These rates are considerably lower than those reported for other rain forest areas in Australia, and are within the lower ranges recorded for other tropical rain forests. Litterfall was found to be strongly seasonal at all sites with the maximum falls occurring from the end of the dry season to the end of the wet season. The between year variability of the sites was quite low, with the ratio of maximum to minimum production ranging from 1.05 to 1.20. The average percentages of leaves, wood and reproductive material in litterfall were similar at each site. Leaves were the dominant component of litterfall with the average proportion of the total litterfall ranging from 72% to 76% over the study period at each of the four sites. At certain times, however, the fall of wood and reproductive material was quite significant, comprising as much as 71% and 34% of litterfall respectively. A strong negative correlation was found between the fall of leaves and wood at all sites.

Mean annual quantities of solar radiation absorbed during various stages of regeneration of a tropical rain forest in the upper Rio Negro valley of Colombia and Venezuela were estimated for the consecutive intervals between clear-cut and 1,3, 10, 20, 35, 60, 80 and 200 years of growth. Forest phytomass and litter fall data from each of these stages were used to calculate the mean annual net dry matter production per unit of absorbed photosynthelically active radiation (PAR), the PAR conversion efficiency. The quantities of PAR absorbed by the forest stands were calculated from the leaf area index values with an extinction coefficient for PAR of 0.74, a PAR albedo of 0.04, and an annual mean incoming PAR of 2.86GJ m-2y-1. Efficiency decreased with increase in successional stage. During the first 10 years of regrowth, the efficiency of conversion of PAR into above-ground phylomass averaged 0.23 g MJ-1, decreasing to 0.07 g MJ-1 over the following 50 years. Inclusion of annual root production in the calculations resulted in a small increase in PAR conversion efficiency; however, efficiency was more than doubled for some periods when the annual leaf and twig lillerfall were included. Efficiency values for above-ground production were much lower than PAR conversion efficiency values estimated for above-ground production of temperate forests.

Production and nutrient content of small litterfall were studied for one year in four plantations of Agathis dammara ranging in age from 7 to 35 years old, in a 12-year-old stand of Pinus merkusii and in an approximately three-year-old shrub community dominated by Eupatorium species. The dense Pinus plantation produced considerably more litter (10 t/ha/yr, with 5.5 t/ha as needles) than any of the Agathis stands (2.6–6.2 t/ha/yr; 2.3–3.5 t/ha as leaves) or nearby Lower Montane Rain Forest (6.8 t/ha/yr; 5.4 t/ha as leaves). Litterfall in the Agathis forests increased with stand age. The shrubs produced sizeable amounts of litter (3.4 t/ha/yr). The various species responded differently to a drought occurring during the study period.

Pine litter generally exhibited low nutrient concentrations (on average 9.2, 1.9, 2.5, 0.4 and 6.0 mg/g of calcium, magnesium, potassium, phosphorus and nitrogen respectively in needle litter), whilst concentrations in shrub litter were generally high (37.8, 4.9, 1.4, 1.05 and 21.5 mg/g for the same elements); Agathis litter usually showed nutrient concentrations that were intermediate between those for pine or shrub litterfall with only minor differences between stands. Comparison of the present findings with observations at other tropical sites suggests that nutrient content of litterfall in plantation forests may approach that of natural forests when the former become about 20 years old. Pine plantations may constitute an exception in this regard.

The demographic history of a flower-cohort of the tropical tree Dicymbe altsonii (Caesalpiniaceae) was studied by collecting litter from flowers, fruits and seeds, and by monitoring seedling survival. Flower production was estimated at 137 to 172 m−2 in a plot of 1 ha. Four percent of the flowers yielded a pod, which contained an average of 2.0 seeds. Post-dispersal mortality over 3 mo was 39% of the initial number of dispersed seeds. Most flowers and fruits were aborted. Insect and vertebrate predation in the tree and on the ground was very low. Reproductive litter production of this species was estimated to be equal to leaf litter production. The amount of phosphorus lost in litter from fruits and flowers was much larger than in leaf litter. This implies that the impact of the biennial flowering events on the nutrient dynamics of the trees and of the ecosystem may be large. It is argued that the reproductive cycle and the pre-dispersal demography of Dicymbe seem to be determined by a scarcity of nutrient resources, and possibly by pollinator limitation.

Litterfall was recorded for one year in Acacia albida woodland in Mana Pools National Park, Zimbabwe. Leaves fell throughout the year, but leaf-fall rate peaked in the wet season. Litterfall was 1.5 t ha−1 year−1, comprising 50% leaves, 20% fruits, 17% ‘fine’ material (mainly frass), 4% flowers and 9% wood and bark. Net above ground production by A. albida trees was 1.7 t ha−1 year−1, including leaf production of 1.0 t ha−1 year−1, twig production of 0.3 t ha-1 year−1 and fruit production of 0.3 t ha−1 year−1. Insects consumed about 26% of leaf production. Net above ground herbaceous production was estimated from rainfall data using regression equations: it was 6.8 t ha−1 year−1 for annual grasses and 2.8 t ha−1 year−1 for forbs. Net above ground primary production in A. albida woodland was 11.3 t ha−1 year−1. The seasonal variations in the nitrogen, phosphorus, potassium, magnesium and calcium concentrations of different A. albida litter components were recorded. The estimated falls of N, P, K, Ca and Mg were 31.1, 2.1, 9.8, 15.0 and 3.4 kg ha−1 year−1 respectively.

An investigation of seasonal changes in litterfall in a dense stand of the invasive tropical shrub Mimosa pigra was carried out in northern Australia between October 1984 and December 1986. Total annual litterfall averaged 758 g m−2, close to the value predicted for the latitude and altitude of the site. Leaf litter formed 56.6% of this total, woody material 22.7%, and reproductive material 20.7%. Total litterfall was strongly seasonal, at a maximum in the late wet season or early dry season, and at a minimum late in the dry season. A broadly similar pattern was shown by the various components of the litter, except for woody litterfall, which followed no strong seasonal trend. Variation in the amount of available soil moisture, as calculated from a simple water balance model, accounted for much of the variation in the components of litterfall.

The proportion of all initiated flower heads which produce seeds was estimated to be between 2.1 and 4.5%. The mean annual seed fall was 9103 m−2, nearly two orders of magnitude higher than most native shrub populations, and it is argued that this is partly a result of the absence of natural enemies in the introduced range.

Litterfall was sampled during one year in two undisturbed Andean forests at altitudes of 2550 and 3370 m in the Colombian Cordillera Central. Total small litterfall at 2550 and 3370 m was 7.03 and 4.31 tha-1 y-1 respectively, of which 4.61 and 2.82 were leaves, 1.06 and 0.76 woody parts, 0.66 and 0.27 reproductive parts, 0.22 and 0.23 epiphytes, 0.47 and 0.23 unclassified.

Clear differences were also found in nutrient concentrations, both between different litter fractions from one site and between equivalent litter fractions from different sites. Weighted mean concentrations for total small litterfall were (mgg-1, at 2550 and 3370m respectively): 11.7 and 7.3 for nitrogen; 0.86 and 0.44 for phosphorus; 8.4 and 3.1 for potassium. Comparison of nutrient concentrations in crown leaves and shed leaves indicated important reallocation of nitrogen (39% at both sites) and phosphorus (45% at 2550m, 65% at 3370m) before leaf shedding.

The results for litterfall amounts, litter nutrient concentrations and reallocation of nutrients in the two forests are consistent with such data obtained in other montane forests in the wet tropics.

The mass of fine litterfall and nutrient circulation through litterfall were determined in four Melrosideros polymorpha/Cibotium spp.-dominated rainforests that differed in substrate age, parent material texture and annual precipitation on Kilauea and Mauna Loa volcanoes on the island of Hawaii. Three of the sites had rates of litterfall of 5.2 Mg ha−1 y−1; the fourth, which was on the most fertile soil, produced 7.0 Mg ha−1 y−1 of litterfall with higher concentrations of nitrogen and phosphorus. Tree ferns of the genus Cibotium cycled relatively large amounts of nitrogen, phosphorus and potassium through litterfall; their contribution to nutrient circulation was disproportionate to their mass in the forest, or in litterfall. The forest on the youngest substrate, which also had the lowest concentrations of nitrogen in litterfall, was fertilized with complete factorial combinations of nitrogen, phosphorus and a treatment consisting of all other plant nutrients. Additions of nitrogen increased the quantity and nitrogen concentration in litterfall during the second year following the initiation of fertilization, while no other treatment had a significant effect. Additions of nitrogen had no effect on litterfall mass or nutrient concentrations in the most nutrient-rich site.