Litterfall Production Along Successional and Altitudinal Gradients of Subtropical Monsoon Evergreen Broadleaved Forests in Guangdong, China

Evaluation of litterfall production is important for understanding nutrient cycling, forest growth, successional pathways, and interactions with environmental variables in forest ecosystems. Litterfall was intensively studied during the period of 1982–2001 in two subtropical monsoon vegetation gradients in the Dinghushan Biosphere Reserve, Guangdong Province, China. The two gradients include: (1) a successional gradient composed of pine forest (PF), mixed pine and broadleaved forest (MF) and monsoon evergreen broadleaved forest (BF), and (2) an altitudinal gradient composed of Baiyunci ravine rain forest (BRF), Qingyunci ravine rain forest (QRF), BF and mountainous evergreen broadleaved forest (MMF). Mean annual litterfall production was 356, 861 and 849 g m⁻² for PF, MF and BF of the successional gradient, and 1016, 1061, 849 and 489 g m⁻² for BRF, QRF, BF and MMF of the altitudinal gradient, respectively. As expected, mean annual litterfall of the pioneer forest PF was the lowest, but rapidly increased over the observation period while those in other forests were relatively stable, confirming that forest litterfall production is closely related to successional stages and growth patterns. Leaf proportions of total litterfall in PF, MF, BF, BRF, QRF and MMF were 76.4%, 68.4%, 56.8%, 55.7%, 57.6% and 69.2%, respectively, which were consistent with the results from studies in other evergreen broadleaved forests. Our analysis on litterfall monthly distributions indicated that litterfall production was much higher during the period of April to September compared to other months for all studied forest types. Although there were significant impacts of some climate variables (maximum and effective temperatures) on litterfall production in some of the studied forests, the mechanisms of how climate factors (temperature and rainfall) interactively affect litterfall await further study.     

西双版纳不同海拔热带雨林凋落量变化研究

在3个海拔梯度(600、1 100和1 600 m)选取8块热带雨林样地,研究了印度季风环境下西双版纳热带季节雨林和山地雨林的凋落动态随海拔的变化及其与气候的关系。在3个梯度上,年平均温度分别为22.1、20.1和16.6 ℃,年平均温度递减率为-0.005 3 ℃·m^-1。随海拔增加,年平均降雨量增加(分别为1 532、1 659和2 011 mm),但旱季的降雨量基本相同(282~295 mm);年蒸发量变化较小(分别为1 369、1 374和1 330 mm);年平均空气相对湿度降低(分别为86%、81%和84%),旱季后期湿度降低更明显;样地土壤含水显著增加。热带季节雨林凋落量(1 072~1 285 g·m^-2·a^-1)显著高于热带山地雨林凋落量(718~1 014 g·m^-2·a^-1)。凋落量和凋落进程变异系数与海拔之间存在线性显著负相关,凋落量与温度线性显著正相关而与降雨量显著负相关。旱季凋落高峰受到空气相对湿度和土壤含水量影响,随海拔增加空气相对湿度降低使得海拔1 105~1 720 m的凋落高峰提前,但土壤含水量继续增加又会使凋落高峰推后。研究结果得出:1)热带季节雨林凋落量与东南亚热带潮湿雨林相近;2)旱季水分限制随海拔增加而变化,影响凋落高峰出现时间;3)随海拔增加,热带山地雨林凋落年进程由季节性向平稳过渡。     

东北地区主要森林生态系统凋落量的比较

以东北地区不同纬度带上4个区域内的典型森林类型为研究对象,采用直接收集法对森林凋落量进行连续3年的观测研究,结果表明,同一地区不同林型间的年凋落量差异显著,凋落量年变化不显著;落叶量占总量的比例最大,在70%以上,落枝和花果皮各占总量的20%以下,落枝量和落叶量之间存在正相关关系,相关系数为0.82;长白山区5种森林类型不同时期内凋落量统计结果显示,秋、冬季的凋落量明显大于春、夏季,前者为后者的2.62～9.14倍.不同凋落组分之间凋落动态不同,落叶以秋季为主,落枝和花果皮则年内分布比较均匀.同一林型的凋落量随纬度增加而减少,并建立了总体森林凋落量和白桦凋落量随纬度变化的数学关系式;总体森林凋落量和白桦林凋落量与温度均呈正相关,相关系数分别为0.95和0.91.     

不同坡位对琉球松人工林凋落物及其养分归还量的影响

对日本冲绳岛北部不同坡位的琉球松人工林凋落物量及其养分归还量进行了为期3年的观测研究,结果表明,琉球松成林年凋落物量为6．54～8．05Mg·hm^-2·年^-1,其凋落高峰出现在6、7月份．台风干扰对年凋落物量及其季节动态影响显著．凋落物不同组分的养分含量差异明显．年均养分归还总量为113．4～154．6kg·hm^-2·年^-1,其中氮素归还量最大,占42．2％;各养分归还量的大小依次为N>Ca>K>Mg>Na>P．两调查林分由于立地条件的显著差异,年均凋落物量及其相应的养分归还量亦存在明显差异．在立地条件较好的下坡,林分P1的年均凋落物量达8．05Mg·hm^-2·年^-1,高于立地条件相对较差的上坡的林分P2的23．1％;其相应的年养分归还量为：N66、04,P1.63,K17．42,Ca48.31,Mg14.65和N。6、57kg·hm^-2·年^-1,分别高于林分P2的39.7％、48.8％、39．4％、32.9％、24.8％和49.3％．两林分凋落物的养分利用效率分别是N为122和138,P为4934和5945,K为462和523,林分P1明显低于林分P2,这与林分P2的立地条件较差相关．可见,所调查的琉球松林是一种高效的养分利用系统。     

沿海拔梯度武夷山3种典型森林凋落物及养分归还动态

为了解武夷山森林凋落物量和养分归还特征,对武夷山3个海拔典型森林的凋落物量和养分动态开展连续3年的监测。结果表明,常绿阔叶林(645 m)、针阔混交林(1 028 m)和针叶林(1 442 m)的凋落物总量分别为471.25、453.77和409.84 g/m~2,森林凋落物总量随海拔升高呈减少的趋势。武夷山3种典型森林的凋落物总量具有明显的季节变化,均呈双峰型,但常绿阔叶林与针阔混交林和针叶林的凋落物总量峰值和次峰值出现时间近似相反。3种典型森林凋落物中落叶占绝对优势(78.1%～87.6%),落枝和其他组分较少。武夷山3种典型森林凋落物的养分年归还量均表现为C N K P,且养分归还总量随海拔升高而减少。常绿阔叶林的凋落物量和养分归还量较高,可能与环境条件和植被特征相关。这为中亚热带森林生态系统养分循环和森林碳循环机理研究提供了理论依据。     

Pantropical trends in mangrove above-ground biomass and annual litterfall

A major paradigm in biosphere ecology is that organic production, carbon turnover and, perhaps, species diversity are highest at tropical latitudes, and decrease toward higher latitudes. To examine these trends in the pantropical mangrove forest vegetation type, we collated and analysed data on above-ground biomass and annual litterfall for these communities. Regressions of biomass and litterfall data show significant relationships with height of the vegetation and latitude. It is suggested that height and latitude are causally related to biomass, while the relationship with litterfall reflects the specific growing conditions at the respective study sites. Comparison of mangrove and upland forest litterfall data shows similar trends with latitude but indicates that mangrove litterfall is higher than upland forest litterfall. The regression equations allow the litterfall/biomass ratio to be simulated, and this suggests that the patterns of organic matter partitioning differ according to latitude.     

雷州半岛桉树人工林凋落物量和养分循环研究

为了解雷州半岛地区桉树人工林凋落物量和养分归还特征,对不同林龄人工林凋落物量和养分动态连续12个月进行监测。结果表明,桉树人工林的凋落物总量为5 a9 a7 a,5 a生桉树人工林的凋落物总量显著高于7和9 a生林分,且7和9 a生林分间没有显著差异。不同林龄桉树人工林凋落物总量具有明显的季节变化,均呈双峰型,峰值出现在雨季初期和末期。林分结构因子同凋落物量相关性不显著,但气候因子中的月均温与凋落物量存在显著正相关关系。不同林龄桉树人工林的养分年循环量由凋落物量和凋落物养分含量共同决定,氮、磷、钾元素的养分年归还总量为9 a7 a5 a,且彼此间达到显著差异。凋落物养分元素归还高峰期集中在雨季前后,以冬季养分归还量较低。因此,为增加桉树人工林产量且利于人工林地力恢复,桉树人工林应适当增加种植年限,且采伐季节选择在冬季。     

浙江建德青冈常绿阔叶林凋落量研究

 本文报道了浙江建德青冈常绿阔叶林的凋落量及各类凋落物的凋落特征。 6年的测定结果表明,青冈林的年均凋落量为5547.6kg/(hm2·a),其中枯叶量占68.32%,枯枝、落果、其它凋落物各占14.82%、15.04%和1.82%。各类凋落量具有明显的季节与年际变化规律,其季节分配还具年际波动现象。各类凋落物的凋落特征从一定角度反映了植物群落一般的生物学与生态学特性,以及植物对特殊环境条件的适应性。     

Is primary production in holm oak forests nutrient limited?

Correlations between primary production and patterns of nutrient use and nutrient availability were investigated in 18 plots in closed holm oak (Quercus ilex L.) stands in the Montseny mountains (NE Spain), searching for evidence of nutrient limitation on primary production. The plots spanned a range of altitudes and slope aspects within a catchment. Nutrients considered were nitrogen (N), phosphorus (P), potassium (K) and magnesium (Mg) in plant samples, and the above plus calcium (Ca) and sodium (Na) in the soil. Primary production was found by summing the annual aboveground biomass increment to the annual litterfall. Across plots, primary production was correlated with the annual return of nutrients in litterfall, but this relationship probably arose from the common effects of the amount of litterfall on both primary production and nutrient return, and not from any nutrient limitation. Primary production was not significantly correlated with nutrient concentrations in mature leaves nor leaf litterfall, nor with absolute or relative foliar retranslocation of nutrients before leaf abscission, nor with the concentration and content (kg/ha) of total N, extractable P, and exchangeable K, Mg, Ca and Na in the upper mineral soil. We conclude that there is no correlational evidence that primary production is nutrient limited in this holm oak forest.     

Litter production,nutrient recycling and litter accumulation inPinus caribaea Morelet var.hondurensis stands in the northern Guniea savanna of Nigeria

Summary The seasonal pattern and quantity of litterfall were studied during a two-year period in two unthinned stands ofPinus caribaea Morelet var. hondurensis Barr. and Golf. in Nigeria. Although pine needles were cast continuously throughout the year, the peak period of litterfall occurred in the dry months of March and April. Mean values of annual litterfall were 3068 and 3665 kg/ha in the two stands aged 7–9 and 9–11 years respectively. Nutrient returns in litterfall in the stands had mean values of 15.0, 0.6, 17.3, 18.2 and 6.3 kg/ha of N, P, K, Ca and Mg respectively. Comparatively low amounts of N and P returned in litterfall were attributed to soil deficiencies of the two elements.Measurements of ground litter showed considerable dry matter accumulation (11378 kg/ha) in the litter layers. Estimates of litter decomposition rate and recycling time showed that it would take 3 to 4 years for the organic matter in annual litterfall to decompose completely as contrasted to about 2 to 5 months often reported under mixed nautral savanna vegetation in the same climatic environment. Similar estimates of nutrient recycling time also showed that between 2 to 4 years were required to mineralize nutrient elements in the annual litterfall; the relative mobilities of the elements were in the order K>Mg>P>NCa.     

福建和溪亚热带雨林凋落物营养元素动态

探讨了福建和溪亚热带雨林主要营养元素通过凋落物的归还动态．结果表明,该森林一年中凋落物不同组分各元素的含量具有波动性,加权平均含量为Ｎ＞Ｋ＞Ｃａ＞Ｍｇ＞Ｐ（落果Ｃａ＝Ｍｇ、落枝Ｃａ＞Ｋ）。通过凋落物归还Ｎ、Ｐ、Ｋ、Ｃａ、Ｍｇ元素的年库流量,分别为８７．４１、３．８２、４０．８０、２８．８１和１５．３２ｋｇｈｍ（－２）,灰分量为２８４．５７ｋｇｈｍ（－２）,其中以落叶形式居最大,分别各占总量的７３．４％、６８．１％、．７７．２％、７２．９％、７４．５％和７９．９％;各元素的年库流量与年凋落物量的月分布相一致,有明显的季节性．森林地表残留物Ｎ、Ｐ、Ｋ、Ｃａ、Ｍｇ元素的累积储量．分别为５４．８３、２．４５、９．６９、１７．０和９．２５ｋｇｈｍ（－２）．五种元素在林下地表的滞留时间,分别为Ｎ０．６１、Ｐ０．６１、Ｋ０．２３、Ｃａ０．５６和Ｍｇ０．５３年．     

Effect of typhoon disturbance on fine litterfall and related nutrient input in a subtropical forest on Okinawa Island, Japan

Typhoons are frequent on Okinawa Island, southwestern Japan. The effects of typhoon disturbance on the patterns of fine litterfall and related nutrient inputs in a subtropical evergreen broad-leaved forest were studied over 5 years from May 1996 to April 2001. Annual fine litterfall averaged 7558 kg ha⁻¹ (range from 6188 to 9439 kg ha⁻¹) for six sampling plots over 5 years, which differed significantly among years (p<0.001) but not among plots (p=0.122). A seasonal maximum was most evident for leaf litter component. Woody litter fell more irregularly through the year, and peak fall varied with typhoon and windstorm. The mean ratio of annual litterfall mass of sexual organs to leaves was 0.06, much lower than that in other tropical and subtropical rain forests. Nutrient concentrations varied in litterfall components, but were not significantly different among plots. The lowest concentrations of N and P in leaf litter were observed in March, which is also the month with the greatest leaf fall. However, the highest concentrations were recorded in typhoon season. Nitrogen and P concentrations were 34% and 106% greater in the green leaves that fell during typhoons than in senescent leaves. Mean nutrient inputs by litterfall were: N 83, P 3.2, K 25, Ca 71, Mg 19, Al 12, Na 10, Fe 0.86 and Mn 3.9 kg ha⁻¹ yr⁻¹, and differed significantly among years for all elements (p<0.0005) and among plots only for K (p<0.05) and Mn (p<0.0001). Typhoon disturbance strongly affected annual fine litterfall and related nutrient inputs, which contributed an average of 30% of the annual litterfall mass, and from 30% to 39% (for different nutrient elements) of annual total nutrient inputs. The results from this study suggest that typhoon-driven maintenance of rapid cycling of P and N and their high availability in soil appears to be an important mechanism to maintain productivity in the subtropical forest on Okinawa Island.     

Temporal and spatial variation of litter production in Sonoran Desert communities

The seasonal pattern of litter production was analyzed in three contiguous desert communities near the southern boundaries of the Sonoran Desert. There was a large spatial variation in annual litter production mainly caused by differences in the composition and structure of vegetation. In the most productive site (Arroyos) annual litterfall was 357 g m^-2yr^-1, a figure higher than some tropical deciduous forests. Litter production was only 60g m^-2yr^-1in the open desert in the plains (Plains) and 157 g m^-2yr^-1 in the thornscrub on the slopes (Hillsides). Topographic and hydrologic features influence the composition, structure and function of the vegetation, modifying the general relationship between rainfall and productivity described for desert ecosystems. The temporal pattern of litter production showed marked seasonality with two main periods of heavy litterfall: one after the summer rains from September to November (autumn litter production) and another after the winter rains from March to May (spring litter production). In the open desert areas, spring litter production was significantly higher than the autumn pulse, while in the slopes, the autumn production was the most important. The Arroyos site produced similar litterfall amounts during the two dry seasons. The species composition defined the season of maximum leaf-fall. In the Plains, the vigorous winter growth of ephemeral and perennial plants made up most of the litter production, while in the Hillsides, most perennials remained dormant throughout the winter-spring period and a significant peak of litterfall occurred only after the summer growth. This difference in growth between seasons was less pronounced in the Arroyos. The timing of maximum production of reproductive and woody litter also differed from site to site.     

Is nutrient availability related to plant nutrient use in humid tropical forests?

Data on soil nutrient availability for humid tropical forests are often reported, but are rarely integrated in an ecologically meaningful way with other measures of nutrient cycling. In this paper, estimates of soil nutrient availability and the inverse of litterfall nutrient concentrations (as an index of plant nutrient use) were compared, using data from 36 sites throughout the humid tropics, to determine if relationships exist between commonly used indices of nutrient cycling for plants and soils. Measures of both extractable and total soil P were significantly and positively correlated with the ratio of litterfall mass/P, particularly for montane tropical forests. Extractable soil P was also significantly correlated with litterfall mass for lowland humid tropical forests, explaining 58% of the variability in litterfall mass. A weak, albeit significant correlation was found between exchangeable soil Ca and litterfall mass/Ca, even though soil extraction techniques vary greatly. No significant relationship was found for total soil N, the most commonly measured soil N pool, and the inverse of litterfall N concentrations. The results suggest that our indices of soil P are related to litterfall processes, but that other measures, particularly total soil N, may not be as relevant to nutrient cycling by the vegetation.     

马占相思 (Acacia mangium)与湿地松 (Pinus elliotii)人工林枯落物层的水文生态功能

分析了马占相思与湿地松人工林枯落物的蓄积量、年凋落量及凋落动态、枯落物层对大气降水的截留、以及枯落物抑制土壤水分蒸发和阻滞径流的效应.结果表明①15龄的马占相思林枯落物蓄积量32.3t/hm2,年凋落量11.14t/hm2,最大持水率253.7%,最大持水量28.26t/hm2;15龄的湿地松林枯落物蓄积量18.7t/hm2,年凋落量7.30t/hm2,最大持水率216.7%,最大持水量15.82hm2;②2种林分对大气降水的截留率分别为15.9%和11.7%,截留率随1次降水降水量(>10mm)的增加而减少;③2～4cm枯落物覆盖下不同含水量的土壤水分蒸发比无覆盖的土壤减少18.2%～78.3%,枯落物层减少土壤水分蒸发的效应随枯落物层厚度和土壤含水量的增大而增加;④2种枯落物对径流流出时间的阻滞效应随径流深(<3mm)和坡度的增加而减小,随枯落物层厚度的增加呈直线增加.通过与部分其它森林类型枯落物层水文生态功能比较,认为马占相思与湿地松林枯落物层具有较为优越的水文生态功能.     

Litter production, leaf litter decomposition and nutrient return in Cunninghamia lanceolata plantations in south China: effect of planting conifers with broadleaved species

This study compared litter production, litter decomposition and nutrient return in pure and mixed species plantations. Dry weight and N, P, K, Ca, Mg quantities in the litterfall were measured in one pure Cunninghamia lanceolata plantation (PC) and two mixed-species plantations of C. lanceolata with Alnus cremastogyne (MCA) and Kalopanax septemlobus (MCK) in subtropical China. Covering 6 years of observations, mean annual litter production of MCA (4.97 Mg·ha⁻¹) and MCK (3.97 Mg·ha⁻¹) was significantly higher than that of PC (3.46 Mg·ha⁻¹). Broadleaved trees contributed 42% of the total litter production in MCA and 31% in MCK. Introduction of broadleaved tree species had no significant effect on litterfall pattern. Total litterfall was greatest in the dry season from November to March. Nutrient returns to the forest floor through leaf litter were significantly higher in both MCA and MCK than in PC (P < 0.05). The amounts of N, K, and Mg returned to the forest floor through leaf litter were highest in the MCA, and P and Ca returns were highest in the MCK. Percent contribution of broadleaf litter to total nutrient returns ranged from 41.7% to 86.9% in MCA and from 49.3% to 74.8% in MCK. The decomposition rate of individual leaf litter increased in the order: C. lanceolata < K. septemlobus < A. cremastogyne. Litter mixing had a positive effect on decomposition rate of the more recalcitrant litter and promoted nutrient return. Relative to mass loss of A. cremastogyne decomposing alone, higher mass loss of the mixture of C. lanceolata and A. cremastogyne was observed after 330 days of decomposition. These results indicate that mixed plantations of different tree species have advantages over monospecific plantations with regards to nutrient fluxes and these advantages have relevance to restoration of degraded sites. Responsible Editor: Alfonso Escudero.     

浙江天童常绿阔叶林演替过程凋落物数量及分解动态

浙江天童常绿阔叶林演替过程森林凋落物量呈增长趋势,其中：乔木叶比例最大,其它器官组分在不同演替阶段的比例不一致,灌木叶和草本、树皮比例随着演替进程而呈减少趋势,树枝和繁殖器官则呈增大趋势。以分解袋法测得凋落物年失重率和指数方程模拟的分解系数Ｋ在演替过程中均呈增长趋势,这与年凋落物数量变化趋势相似。可见,随着演替进程,生物循环加快,群落自肥能力增强。     

浙江天童常绿阔叶演替过程凋落物数量及分解动态

 浙江天童常绿阔叶林演替过程森林凋落物量呈增长趋势,其中：乔木叶比例最大,其它器官组分在不同演替阶段的比例不一致,灌木叶和草本、树皮比例随着演替进程而呈减少趋势,树枝和繁殖器官则呈增大趋势。以分解袋法测得凋落物年失重率和指数方程模拟的分解系数K在演替过程中均呈增长趋势,这与年凋落物数量变化趋势相似。可见,随着演替进程,生物循环加快,群落自肥能力增强。     

贡嘎山峨眉冷杉林凋落物的特征

 研究了贡嘎山峨眉冷杉（Abies fabri）林凋落物量、凋落物特征及其N、P、K、有机碳的归还量,并探讨了它们在演替进程中的变化。结果表明,峨眉冷杉成熟林、中龄林和幼龄林的凋落物量依次为：2 809.925、2 787.086、2 043.585 kg·hm-2·a-1;其中N、P、K的归还量依次为：34.850、33.917、42.571 kg·hm-2· a-1。峨眉冷杉林凋落物量及其N、P、K的归还量较少,与我国其它森林区针叶林存在一定差异,而与美国西北部冷杉林相近。     

Litterfall and Leaf Consumption by Animals in Humid Seasonal Rainforest in Xishuangbanna,China

Litterfall and element return dynamic and leaf consumption by animals were studied in humid seasonal rainforest in Menglun, Xishuangbanna, Southwesten China. 1. The litterfall was l1.29+l.96t/ha·a, and the amount of litterfall component was (t/ha·a±S. E.): for leaves, 7.41±0.82; wood (≤2 cm diameter), 2.16±1.78; flowers, 0.223±0.13; fruits, 0.7244±0.456 and mixed matter (includs plant trash, wastes of animals and died animal bodies from canopy), 0.77±0.18. A marked peak of litterfall occurred during a March-April period of the late dry season. 2. Large wood litterfall (>2 cm diameter) was measured twice during rainy season and dry season, the large wood litterfall mean was 0.82t/ha·a. 3. Returns (kg/ha·a) of the elements to the forest floor in the litterfall were ranged Ca (146.4) > N (57.3) > Mg (39.0) 3> K(31.2) > P (6.33) > Mn (5.49) 2> A1(4.40) > Fe(1.92) > Zn(0.56) > Cu (0.24).and a marked peak of element return coincided with the litterfall's. 4. The leaf area eaten by animals accounted for 16.2% of the total leaf area.