The influence of macro‐litterfall and forest structure on litterfall damage to seedlings

Abstract Microdisturbance to seedlings is important because it can differentially affect the mortality and recruitment of seedlings of forest tree species and thereby ultimately affect community composition. Microdisturbance due to litterfall has been shown to vary greatly in its influence on seedling survival among and within forests, and yet there have been no previous studies that investigate the cause of these differences. In this study the influence of macro‐litterfall on seedling damage is investigated in five complex temperate forests in New Zealand. Litterfall damage to artificial seedlings in these forests was strongly correlated with macro‐leaf‐fall (leaves > 30 cm × 1.5 cm) dry weight and total macro‐litterfall (leaves and deadwood > 30 cm × 1.5 cm) surface area (R² = 0.99, P < 0.005 for each). Protective vegetation within 2 m of the ground (mostly lianes and woody shrubs) reduced the risk of litterfall damage by up to 84%. Hitherto unexplained differences in litterfall damage to seedlings found among, and within, forests (tropical and temperate) may therefore be due to differences in rates of macro‐leaf‐fall and forest structure. These results are important because they suggest that subtle differences in forest structure, and species composition, may influence regeneration patterns through the litterfall microdisturbance regime.     

Detritus Production and Soil N Transformations in Old-Growth Eastern Hemlock and Sugar Maple Stands

To examine the linkage between forest cover type, litter inputs, and patterns of net N mineralization versus the turnover of N among soil microbes, we measured both the net and gross rates of N mineralization in replicated, adjacent old-growth eastern hemlock [Tsuga canadensis(L.) Carr.] or sugar maple (Acer saccharum Marsh.) stands in upper Michigan. Mean aboveground net primary production and annual litterfall mass were significantly higher (P < 0.01) in the maple forests (870 g·m^-2·y^-1 and 439 g·m^-2·y^-1, respectively) than in the hemlock forests (480 g·m^-2·y^-1 and 344 g·m^-2·y^-1, respectively). Forest floor and coarse woody debris mass, however, were significantly lower (P < 0.05) in the maple forests (2.2 and 0.1 kg·m^-2, respectively) than in the hemlock forests (2.9 and 0.2 kg·m^-2, respectively). Litterfall N concentration was not significantly different (P > 0.10) between the two forest types. In situ gross rates of N mineralization were higher (P < 0.06) in the maple forests than in the hemlock forests (7.5 and 6.1 mg N·kg soil^-1·d^-1 respectively), but in situ net N mineralization varied independently of forest type and stand-level litterfall N concentration. Cover type–dependent differences in detritus production and detritus C quality appear to result in different N turnover rates, but the balance between gross mineralization and immobilization of N is very sensitive to within stand variability and varies at a scale smaller than cover type alone can predict. Received 3 Feburary 1999; accepted 27 August 1999.     

Original Articles: Nitrogen Mineralization in Two Unpolluted Old-Growth Forests of Contrasting Biodiversity and Dynamics

Studies in unpolluted, old-growth forests in the coastal range of southern Chile (42°30′S) can provide a baseline for understanding how forest ecosystems are changing due to the acceleration of nitrogen (N) inputs that has taken place over the last century. Chilean temperate forests, in contrast to their northern hemisphere counterparts, exhibit extremely low losses of inorganic N to stream waters. The objectives of this study were (a) to determine whether low inorganic N outputs in these forests were due to low rates of N mineralization or nitrification, and (b) to examine how biodiversity (defined as number of dominant tree species) and forest structure influence N mineralization and overall patterns of N cycling. Studies were conducted in a species-poor, conifer-dominated (Fitzroya cupressoides) forest with an even-aged canopy, and in a mixed-angiosperm (Nothofagus nitida) forest with a floristically more diverse and unstable canopy. Nitrogen mineralization rates measured in laboratory assays varied seasonally, reaching 6.0 μg N/g DW/day in both forests during late summer. Higher values were related to higher microbial activity, larger pools of labile inorganic N, and increased fine litter inputs. Field assays, conducted monthly, indicated positive net flux from N mineralization mainly from December to January in both forests. Annual net flux of N from mineralization varied from 20 to 23 kg/ha/year for the Fitzroya forest and from 31 to 37 kg/ha/year for the Nothofagus forest. Despite low losses of inorganic N to streams, N mineralization and nitrification are not inhibited in these forests, implying the existence of strong sinks for NO₃ ⁻ in the ecosystem. Field N mineralization rates were two times higher in the Nothofagus forest than in the Fitzroya forest, and correlated with greater N input via litterfall, slightly higher soil pH, and narrower carbon (C)–nitrogen ratios of soils and litter in the former. Differences in N mineralization between the two forest types are attributed to differences in biotic structure, stand dynamics, and site factors. Median values of net N mineralization rates in these southern hemisphere forests were lower than median rates for forests in industrialized regions of North America, such as the eastern and central USA. We suggest that these high N mineralization rates may be a consequence of enhanced atmospheric N deposition.     

Soil Nutrients Limit Fine Litter Production and Tree Growth in Mature Lowland Forest of Southwestern Borneo

Efforts to improve models of terrestrial productivity and to understand the function of tropical forests in global carbon cycles require a mechanistic understanding of spatial variation in aboveground net primary productivity (ANPP) across tropical landscapes. To help derive such an understanding for Borneo, we monitored aboveground fine litterfall, woody biomass increment and ANPP (their sum) in mature forest over 29 months across a soil nutrient gradient in southwestern Kalimantan. In 30 (0.07 ha) plots stratified throughout the watershed (∼340 ha, 8–190 m a.s.l.), we measured productivity and tested its relationship with 27 soil parameters. ANPP across the study area was among the highest reported for mature lowland tropical forests. Aboveground fine litterfall ranged from 5.1 to 11.0 Mg ha⁻¹ year⁻¹ and averaged 7.7 ± 0.4 (mean ± 95 C.I.). Woody biomass increment ranged from 5.8 to 23.6 Mg ha⁻¹ year⁻¹ and averaged 12.0 ± 2.0. Growth of large trees (≥60 cm dbh) contributed 38–82% of plot-wide biomass increment and explained 92% of variation among plots. ANPP, the sum of these parameters, ranged from 11.1 to 32.3 Mg ha⁻¹ year⁻¹ and averaged 19.7 ± 2.2. ANPP was weakly related to fine litterfall (r ² = 0.176), but strongly related to growth of large trees at least 60 cm dbh (r ² = 0.848). Adjusted ANPP after accounting for apparent “mature forest bias” in our sampling method was 17.5 ± 1.2 Mg ha⁻¹ year⁻¹.Relating productivity measures to soil parameters showed that spatial patterning in productivity was significantly related to soil nutrients, especially phosphorus (P). Fine litterfall increased strongly with extractable P (r ² = 0.646), but reached an asymptote at moderate P levels, whereas biomass increment (r ² = 0.473) and ANPP (r ² = 0.603) increased linearly across the gradient. Biomass increment of large trees was more frequently and strongly related to nutrients than small trees, suggesting size dependency of tree growth on nutrients. Multiple linear regression confirmed the leading importance of soil P, and identified Ca as a potential co-limiting factor. Our findings strongly suggest that (1) soil nutrients, especially P, limit aboveground productivity in lowland Bornean forests, and (2) these forests play an important, but changing role in carbon cycles, as canopy tree logging alters these terrestrial carbon sinks. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effect of different anthropogenic disturbances on litterfall of a dominant pioneer rain forest tree in Gabon

Tropical rain forests worldwide are affected by anthropogenic disturbances, and secondary forests that develop afterwards are often dominated by pioneer tree species, but the consequences of different anthropogenic disturbances on nutrient cycling are poorly understood. Because most nutrient cycling in tropical rain forests occurs through litterfall and in the soil organic layer, we measured litterfall of a widespread and dominant pioneer tree, okoume (Aucoumea klaineana, Burseraceae) in Gabon, in one forest previously disturbed by logging and in another by agriculture. Litterfall of okoume trees, measured over 14 months, was 18.2 Mg ha⁻¹ year⁻¹ in the formerly logged forest, which was 72.6% greater than in the forest more recently disturbed by agriculture (10.6 Mg ha⁻¹ year⁻¹). Litter decomposition rates were more rapid in the formerly logged forest, and this may explain why litter thickness was 32% lower in the formerly logged forest, despite the greater litterfall within it. Our results highlight that two widespread anthropogenic disturbances in Gabonese rain forests have significantly different consequences on litterfall of a dominant tree and therefore are likely to have different effects on nutrient cycling and forest ecosystem function.     

Non‐symbiotic nitrogen fixation during leaf litter decomposition in an old‐growth temperate rain forest of Chiloé Island,southern Chile: Effects of single versus mixed species litter

Heterotrophic nitrogen fixation is a key ecosystem process in unpolluted, temperate old‐growth forests of southern South America as a source of new nitrogen to ecosystems. Decomposing leaf litter is an energy‐rich substrate that favours the occurrence of this energy demanding process. Following the niche ‘complementarity hypothesis’, we expected that decomposing leaf litter of a single tree species would support lower rates of non‐symbiotic N fixation than mixed species litter taken from the forest floor. To test this hypothesis we measured acetylene reduction activity in the decomposing monospecific litter of three evergreen tree species (litter C/N ratios, 50–79) in an old‐growth rain forest of Chiloé Island, southern Chile. Results showed a significant effect of species and month (anova , Tukey's test, P < 0.05) on decomposition and acetylene reduction rates (ARR), and a species effect on C/N ratios and initial % N of decomposing leaf litter. The lowest litter quality was that of Nothofagus nitida (C/N ratio = 78.7, lignin % = 59.27 ± 4.09), which resulted in higher rates of acetylene reduction activity (mean = 34.09 ± SE = 10.34 nmol h^?1 g^?1) and a higher decomposition rate (k = 0.47) than Podocarpus nubigena (C/N = 54.4, lignin % = 40.31 ± 6.86, Mean ARR = 4.11 ± 0.71 nmol h^?1 g^?1, k = 0.29), and Drimys winteri (C/N = 50.6, lignin % = 45.49 ± 6.28, ARR = 10.2 ± 4.01 nmol h^?1 g^?1, k = 0.29), and mixed species litter (C/N = 60.7, ARR = 8.89 ± 2.13 nmol h^?1g^?1). We interpret these results as follows: in N‐poor litter and high lignin content of leaves (e.g. N. nitida) free‐living N fixers would be at competitive advantage over non‐fixers, thereby becoming more active. Lower ARR in mixed litter can be a consequence of a lower litter C/N ratio compared with single species litter. We also found a strong coupling between in situ acetylene reduction and net N mineralization in surface soils, suggesting that as soon N is fixed by diazotroph bacteria it may be immediately incorporated into mineral soil by N mineralizers, thus reducing N immobilization.     

Nitrogen Oxide Fluxes and Nitrogen Cycling during Postagricultural Succession and Forest Fertilization in the Humid Tropics

The effects of changes in tropical land use on soil emissions of nitrous oxide (N₂O) and nitric oxide (NO) are not well understood. We examined emissions of N₂O and NO and their relationships to land use and forest composition, litterfall, soil nitrogen (N) pools and turnover, soil moisture, and patterns of carbon (C) cycling in a lower montane, subtropical wet region of Puerto Rico. Fluxes of N₂O and NO were measured monthly for over 1 year in old (more than 60 years old) pastures, early- and mid-successional forests previously in pasture, and late-successional forests not known to have been in pasture within the tabonuco (Dacryodes excelsa) forest zone. Additional, though less frequent, measures were also made in an experimentally fertilized tabonuco forest. N₂O fluxes exceeded NO fluxes at all sites, reflecting the consistently wet environment. The fertilized forest had the highest N oxide emissions (22.0 kg N · ha⁻¹· y⁻¹). Among the unfertilized sites, the expected pattern of increasing emissions with stand age did not occur in all cases. The mid-successional forest most dominated by leguminous trees had the highest emissions (9.0 kg N · ha⁻¹· y⁻¹), whereas the mid-successional forest lacking legumes had the lowest emissions (0.09 kg N · ha⁻¹· y⁻¹). N oxide fluxes from late-successional forests were higher than fluxes from pastures. Annual N oxide fluxes correlated positively to leaf litter N, net nitrification, potential nitrification, soil nitrate, and net N mineralization and negatively to leaf litter C:N ratio. Soil ammonium was not related to N oxide emissions. Forests with lower fluxes of N oxides had higher rates of C mineralization than sites with higher N oxide emissions. We conclude that (a) N oxide fluxes were substantial where the availability of inorganic N exceeded the requirements of competing biota; (b) species composition resulting from historical land use or varying successional dynamics played an important role in determining N availability; and (c) the established ecosystem models that predict N oxide loss from positive relationships with soil ammonium may need to be modified. Received 22 February 2000; accepted 6 September 2000.     

Litterfall and Element Fluxes in a Natural Hardwood Forest and a Chinese‐fir Plantation Experiencing Frequent Typhoon Disturbance in Central Taiwan

Chinese fir (Cunninghamia lanceolata) is the most important forest plantation species in subtropical Asia and is rapidly replacing natural forests. Such land‐use change may affect ecosystem nutrient cycling through changes in litterfall nutrient flux. Tropical cyclones often cause pulses of litterfall. Previous studies, however, have mostly focused on the effects of a single cyclone with little effort examining the effects of repeated cyclones. We examined litterfall in a natural hardwood forest and a Chinese‐fir plantation in central Taiwan experiencing an average of one typhoon per year. The natural hardwood forest had 54 percent higher annual litterfall (11,400 kg/ha/yr) than the Chinese‐fir plantation (7400 kg/ha/yr). Four typhoon‐affected months (typhoon period) contributed to approximately 60 percent of the litterfall and litterfall element flux in the natural hardwood forest and 80 percent in the Chinese‐fir plantation, with contributions from individual typhoons varied by more than twofold. Litterfall N and P concentrations were significantly higher in typhoon period than in non‐typhoon period, likely the result of limited retranslocation. Precipitation was a better predictor of quantity of typhoon‐associated litterfall than wind velocity. Both types of forests in southeastern China beyond the reach of typhoons have litterfall peaks in the dry season. In contrast, we measured higher litterfall during the typhoon period than during the dry season, suggesting that in regions with frequent cyclones, cyclones drive temporal variation of litterfall. Global climate change is affecting the frequency and intensity of cyclones; therefore, knowledge of typhoon‐litterfall dynamics is indispensable for understanding the effects of climate change on ecosystem nutrient cycling.     

神农架海拔梯度上4种典型森林凋落物现存量及其养分循环动态

研究神农架地区典型森林凋落物现存量及其养分动态对认识我国北亚热带森林生态系统养分循环过程及森林碳循环的机理具有重要的参考价值。通过对神农架海拔梯度上4种典型森林常绿阔叶林、常绿落叶阔叶混交林、落叶阔叶林及亚高山针叶林凋落物年凋落量及其养分归还量的研究,发现:森林凋落物量随海拔增加呈现先上升后降低的趋势,由低海拔到高海拔,凋落物年凋落量分别为6807.97、7118.14、6975.2和4250.67 kg/hm²。各森林类型凋落物量年变化呈双峰型,高峰期出现在4-5月份、11月份。凋落物养分归还以N 最高(132.06、162.29、157.12和185.77 kg/hm²),以P最少(4.62、4.39、8.24和4.15 kg/hm²),养分归还总量随海拔高度增加而减少。     

Biological Nitrogen Fixation in Two Tropical Forests: Ecosystem-Level Patterns and Effects of Nitrogen Fertilization

Humid tropical forests are often characterized by large nitrogen (N) pools, and are known to have large potential N losses. Although rarely measured, tropical forests likely maintain considerable biological N fixation (BNF) to balance N losses. We estimated inputs of N via BNF by free-living microbes for two tropical forests in Puerto Rico, and assessed the response to increased N availability using an on-going N fertilization experiment. Nitrogenase activity was measured across forest strata, including the soil, forest floor, mosses, canopy epiphylls, and lichens using acetylene (C₂H₂) reduction assays. BNF varied significantly among ecosystem compartments in both forests. Mosses had the highest rates of nitrogenase activity per gram of sample, with 11 ± 6 nmol C₂H₂ reduced/g dry weight/h (mean ± SE) in a lower elevation forest, and 6 ± 1 nmol C₂H₂/g/h in an upper elevation forest. We calculated potential N fluxes via BNF to each forest compartment using surveys of standing stocks. Soils and mosses provided the largest potential inputs of N via BNF to these ecosystems. Summing all components, total background BNF inputs were 120 ± 29 μg N/m²/h in the lower elevation forest, and 95 ± 15 μg N/m²/h in the upper elevation forest, with added N significantly suppressing BNF in soils and forest floor. Moisture content was significantly positively correlated with BNF rates for soils and the forest floor. We conclude that BNF is an active biological process across forest strata for these tropical forests, and is likely to be sensitive to increases in N deposition in tropical regions.     

Conversion of secondary broadleaved forest into Chinese fir plantation alters litter production and potential nutrient returns

Litter production, litter standing crop, and potential nutrient return via litterfall to soil were studied during a 4-year period (January 2004–December 2007) in a Chinese fir (Cunninghamia lanceolata (Lamb.) Hook) plantation and a secondary broadleaved forest in Hunan Province in subtropical China. Mean annual litterfall in the sampling sites varied from 358 g m⁻² in the pure plantation to 669 g m⁻² in the secondary broadleaved forest. Total litterfall followed a bimodal distribution pattern for both forests. Amount of litterfall was also related to the air temperature in both forests. During the period under this study, annual variation in the total litterfall in the pure plantation was significantly higher than that in the secondary broadleaved forest. Litterfall was markedly seasonal in the both forests. Leaf proportions of litterfall in the pure plantation and secondary broadleaved forest were 58.1 and 61.7%, respectively. Total potential nutrient returns to the soil through litterfall in the pure plantation were only 46.2% of those in the secondary broadleaved forest. Total litter standing crop was 913 and 807 g m⁻² in the pure plantation and secondary broadleaved forest, respectively. Our results confirm that conversion from a secondary broadleaved forest into a pure coniferous plantation changes the functioning of the litter system.     

Decline of soil nitrogen mineralization and nitrification during forest conversion of evergreen broad-leaved forest to plantations in the subtropical area of Eastern China

We examined soil nitrogen (N) mineralization and nitrification rates, and soil and forest floor properties in one native forest: evergreen broad-leaved forest (EBLF), one secondary shrubs (SS), and three adjacent plantation forests: Chinese fir plantation (CFP), bamboo plantation (BP) and waxberry groves (WG) in Tiantong National Forest Park, Eastern China. All forests showed seasonal dynamics of N mineralization and nitrification rates. Soil N mineralization rate was highest in EBLF (1.6 ± 0.3 mg-N kg⁻¹ yr⁻¹) and lowest in CFP (0.4 ± 0.1 mg-N kg⁻¹ yr⁻¹). Soil nitrification rate was also highest in EBLF (0.6 ± 0.1 mg-N kg⁻¹ yr⁻¹), but lowest in SS (0.02 ± 0.01 mg-N kg⁻¹ yr⁻¹). During forest conversion of EBLF to SS, CFP, BP and WG, soil N mineralization rate (10.7%, 73%, 40.3% and 69.8%, respectively), soil nitrification rate (94.9%, 32.2%, 33.9% and 39%, respectively), and soil N concentration (50%, 65.4%, 78.9% and 51.9%, respectively) declined significantly. Annual soil N mineralization was positively correlated with total C and N concentrations of surface soil and total N concentration of forest floor, and negatively correlated with soil bulk density, soil pH and C:N ratio of forest floor across the five forests. Annual soil nitrification was positively correlated with total C concentration of surface soil and N concentration of forest floor, and negatively correlated with soil bulk density and forest floor mass. In contrast, annual soil nitrification was not correlated to pH value, total N concentration, C:N ratio of surface soil and total C concentration and C:N ratio of forest floor.     

Emissions of nitrous oxide from three tropical forests in Southern China in response to simulated nitrogen deposition

Emissions of nitrous oxide (N₂O) from the soil following simulated nitrogen (N) deposition in a disturbed (pine), a rehabilitated (pine and broadleaf mixed) and a mature (monsoon evergreen broadleaf) tropical forest in southern China were studied. The following hypotheses were tested: (1) addition of N will increase soil N₂O emission in tropical forests; and (2) any observed increase will be more pronounced in the mature forest than in the disturbed or rehabilitated forest due to the relatively high initial soil N concentration in the mature forest. The experiment was designed with four N treatment levels (three replicates; 0, 50, 100, 150 kg N ha⁻¹ year⁻¹ for C (Control), LN (Low-N), MN (Medium-N), and HN (High-N) treatment, respectively) in the mature forest, but only three levels in the disturbed and rehabilitated forests (C, LN and MN). Between October 2005 to September 2006, soil N₂O flux was measured using static chamber and gas chromatography methodology. Nitrogen had been applied previously to the plots since July 2003 and continued during soil N₂O flux measurement period. The annual mean rates of soil N₂O emission in the C plots were 24.1 ± 1.5, 26.2 ± 1.4, and 29.3 ± 1.6 μg N₂O–N m⁻² h⁻¹ in the disturbed, rehabilitated and mature forest, respectively. There was a significant increase in soil N₂O emission following N additions in the mature forest (38%, 41%, and 58% when compared to the C plots for the LN, MN, and HN plots, respectively). In the disturbed forest a significant increase (35%) was observed in the MN plots, but not in the LN plots. The rehabilitated forest showed no significant response to N additions. Increases in soil N₂O emission occurred primarily in the cool-dry season (November, December and January). Our results suggest that the response of soil N₂O emission to N deposition in tropical forests in southern China may vary depending on the soil N status and land-use history of the forest.     

Impacts of forest management type and season on soil carbon fluxes in Eastern Mau Forest,Kenya

The value of ecosystems functions performed by forests in the climate change era has prompted increasing attention towards assessment of carbon stocks and fluxes in tropical forests. The aim of this study was to understand how forest management approaches and environmental controls impacted on soil CO₂ efflux in a tropical Eastern Mau forest which is one of the blocks of the greater Mau complex in Kenya. Nested experimental design approach was employed where 32 plots were nested into four blocks (disturbed natural, undisturbed natural, plantation and glades). In 10 m² plots, data were collected on soil CO₂ efflux, soil temperature and soil moisture using soda lime methods, direct measurement and proxy techniques, respectively. There was significant forest management type effect (F_3,127 = 3.01, p = 0.033) and seasonality effect (t test = 3.31, df = 1, p < 0.05) on mean soil CO₂ efflux. The recorded mean soil CO₂ efflux levels were as follows: plantation forest (9.219 ± 3.067 g C M^?2 day^?1), undisturbed natural forest (8.665 ± 4.818 g C M^?2 day^?1), glades (8.592 ± 3.253 g C M^?2 day^?1) and disturbed natural forest (7.198 ± 3.457 g C M^?2 day^?1). The study concludes that managing a forest in plantation form is primarily responsible for forest soil CO₂ efflux levels due to aspects such as increased microbial activity and root respiration. However, further studies are required to understand the role and impact of soil CO₂ efflux on the greater forest carbon budget.     

长期氮沉降下杉木人工林凋落物与土壤的C、N、P化学计量特征

为研究长期氮沉降条件下林木凋落物与土壤养分之间的关系,该文以亚热带杉木(Cunninghamia lanceolata)人工林为研究对象,分析了模拟氮沉降处理第12年时杉木林凋落物不同组分(叶、枝、果)与不同土层土壤(0～20 cm、20～40 cm、40～60 cm)的C、N、P含量及其化学计量比。氮沉降处理分4个水平,分别为N0(0 kg N·hm^-2·a^-1)、N1(60 kg N·hm^-2·a^-1)、N2(120 kg N·hm^-2·a^-1)、N3(240 kg N·hm^-2·a^-1),每处理重复3次。结果表明:(1)凋落物各组分的C、N、P含量及其化学计量比均高于土壤; 凋落物和土壤化学计量比均表现为C/P>C/N>N/P; 凋落物不同组分的C、N含量表现为叶>果>枝,而P含量表现为叶>枝>果。(2)12 a氮沉降增加了凋落物叶、枝和果的N含量,增幅分别为4.24%、15.97%、6.47%; 同时增加了凋落物枝N/P,降低了凋落物枝C含量、C/N和C/P; 中-高氮沉降(N2、N3)增加了土壤N含量,低氮沉降(N1)增加了土壤C/P、N/P。(3)相关性分析表明凋落物N与土壤N显著正相关,土壤C/P与凋落物C/P、N/P显著负相关,土壤P与凋落物N/P显著负相关。综上结果说明凋落物N是土壤N的重要N素来源之一,而土壤N可能是决定长期氮沉降后凋落物N/P的主要因素。     

Litterfall and nutrient dynamics shift in tropical forest restoration sites after a decade of recovery

Multi‐year studies comparing changes in litterfall biomass and nutrient inputs in sites under different restoration practices are lacking. We evaluated litterfall dynamics and nutrient inputs at 5 yr and after a decade of recovery in four treatments (natural regeneration—no planting, plantation—entire area planted, tree islands—planting in patches, and reference forest) at multiple sites in an agricultural landscape in southern Costa Rica. We inter‐planted two native species (Terminalia amazonia and Vochysia guatemalensis) and two naturalized N‐fixing species (Inga edulis and Erythrina poeppigiana) in plantation and island treatments. Although litterfall N was higher in plantations in the first sampling period, litter production and overall inputs of C, N, Ca, Mg, P, Cu, Mn, and Fe did not differ between island, plantation, or reference forest after a decade; however, all were greater than in natural regeneration. Potassium inputs were lower in the natural regeneration, intermediate in island and plantation, and greater in reference forest. The percentage of litterfall comprised by the N‐fixing planted species declined by nearly two‐thirds in both plantations and islands between sampling periods, while the percentage of V. guatemalensis more than doubled, and the percentage from naturally regenerated species increased from 27 to 47 percent in islands. Island and plantation treatments were equally effective at restoring litterfall and nutrient inputs to levels similar to the reference system. The nutrient input changed substantially over the 7‐yr interval between measurements, reflecting shifts in vegetation composition and demonstrating how rapidly nutrient cycling dynamics can change in recovering forests.     

Effects of nitrogen additions on above- and belowground carbon dynamics in two tropical forests

Anthropogenic nitrogen (N) deposition is increasing rapidly in tropical regions, adding N to ecosystems that often have high background N availability. Tropical forests play an important role in the global carbon (C) cycle, yet the effects of N deposition on C cycling in these ecosystems are poorly understood. We used a field N-fertilization experiment in lower and upper elevation tropical rain forests in Puerto Rico to explore the responses of above- and belowground C pools to N addition. As expected, tree stem growth and litterfall productivity did not respond to N fertilization in either of these N-rich forests, indicating a lack of N limitation to net primary productivity (NPP). In contrast, soil C concentrations increased significantly with N fertilization in both forests, leading to larger C stocks in fertilized plots. However, different soil C pools responded to N fertilization differently. Labile (low density) soil C fractions and live fine roots declined with fertilization, while mineral-associated soil C increased in both forests. Decreased soil CO₂ fluxes in fertilized plots were correlated with smaller labile soil C pools in the lower elevation forest (R² = 0.65, p < 0.05), and with lower live fine root biomass in the upper elevation forest (R² = 0.90, p < 0.05). Our results indicate that soil C storage is sensitive to N deposition in tropical forests, even where plant productivity is not N-limited. The mineral-associated soil C pool has the potential to respond relatively quickly to N additions, and can drive increases in bulk soil C stocks in tropical forests.     

中亚热带4种森林凋落物量、组成、动态及其周转期

为研究亚热带次生林保护对森林生态系统养分循环等功能过程的影响。采用凋落物直接收集法,比较湘中丘陵区3种次生林(马尾松+石栎针阔混交林、南酸枣落叶阔叶林、石栎+青冈常绿阔叶林)和杉木人工林的凋落物量、组成特征及其周转期。结果表明:4种林分年凋落物量在414.4—818.2 g m-2a-1之间,3种次生林显著高于杉木人工林,3种次生林两两之间差异不显著,落叶对林分凋落物量的贡献最大,占林分凋落物量的59.9%—66.6%。杉木人工林和南酸枣落叶阔叶林的凋落物量月动态变化呈"双峰型",马尾松+石栎针阔混交林、石栎+青冈常绿阔叶林呈"不规则型"。优势树种的凋落物量对其林分凋落物量的贡献随林分树种多样性的增加而下降,杉木、马尾松凋落物量的月动态与其林分凋落物量的月动态基本呈一致变化趋势,但南酸枣、青冈、石栎没有一致的变化趋势。杉木人工林凋落物分解率最低(0.31),周转期最长(3.2 a),南酸枣落叶阔叶林分解率最高(0.45),周转期最低(2.2 a),凋落物的分解速率和周转随林分树种多样性增加而加快。可见,次生林凋落物量大,且分解快,周转期短,有利于养分归还和具有良好地力维持的能力。     

Spatial Variability in Litterfall,Litter Standing Crop and Litter Quality in a Tropical Rain Forest Region

Understanding the spatial variability in plant litter processes is essential for accurate comprehension of biogeochemical cycles and ecosystem function. We assessed spatial patterns in litter processes from local to regional scales, at sites throughout the wet tropical rain forests of northern Australia. We aimed to determine the controls (e.g., climate, soil, plant community composition) on annual litter standing crop, annual litterfall rate and in situ leaf litter decomposability. The level of spatial variance in these components, and leaf litter N, P, Ca, lignin, α‐cellulose and total phenolics, was determined from within the scale of subregion, to site (1 km transects) to local/plot (~30 m²). Overall, standing crop was modeled with litterfall and its chemical composition, in situ decomposability, soil Na, and topography (r² = 0.69, 36 plots). Litterfall was most closely aligned with plant species richness and stem density (negative correlation); leaf decomposability with leaf‐P and lignin, soil Na, and dry season moisture (r² = 0.89, 40 plots). The predominant scale of variability in litterfall rates was local (plot), while litter standing crop and α‐cellulose variability was more evenly distributed across spatial scales. Litter decomposability, N, P and phenolics were more aligned with subregional differences. Leaf litter C, lignin and Ca varied most at the site level, suggesting more local controls. We show that variability in litter quality and decomposability are more easily accounted for spatially than litterfall rates, which vary widely over short distances possibly in response to idiosyncratic patterns of disturbance.     

Complex effects of mammalian grazing on extramatrical mycelial biomass in the Scandes forest‐tundra ecotone

Mycorrhizal associations are widespread in high‐latitude ecosystems and are potentially of great importance for global carbon dynamics. Although large herbivores play a key part in shaping subarctic plant communities, their impact on mycorrhizal dynamics is largely unknown. We measured extramatrical mycelial (EMM) biomass during one growing season in 16‐year‐old herbivore exclosures and unenclosed control plots (ambient), at three mountain birch forests and two shrub heath sites, in the Scandes forest‐tundra ecotone. We also used high‐throughput amplicon sequencing for taxonomic identification to investigate differences in fungal species composition. At the birch forest sites, EMM biomass was significantly higher in exclosures (1.36 ± 0.43 g C/m²) than in ambient conditions (0.66 ± 0.17 g C/m²) and was positively influenced by soil thawing degree‐days. At the shrub heath sites, there was no significant effect on EMM biomass (exclosures: 0.72 ± 0.09 g C/m²; ambient plots: 1.43 ± 0.94). However, EMM biomass was negatively related to Betula nana abundance, which was greater in exclosures, suggesting that grazing affected EMM biomass positively. We found no significant treatment effects on fungal diversity but the most abundant ectomycorrhizal lineage/cortinarius, showed a near‐significant positive effect of herbivore exclusion (p = .08), indicating that herbivory also affects fungal community composition. These results suggest that herbivory can influence fungal biomass in highly context‐dependent ways in subarctic ecosystems. Considering the importance of root‐associated fungi for ecosystem carbon balance, these findings could have far‐reaching implications.