Spatial and temporal variability of canopy structure in a tropical moist forest

Tree fall gaps are widely considered to play a prominent role in the maintenance of species diversity, while the spatiotemporal variability of canopy structure within closed forest stands is largely ignored. In this study we examined the vertical and horizontal components of canopy structure and its seasonal variability in a tropical wet semideciduous rainforest in Panama. Leaf area indices (LAI) were derived from measurements of diffuse radiation and empirically-based leaf angle distribution by mathematical inversion of a light interception model. Vertical distribution of LAI was non-homogeneous with 50% of the leaf area being concentrated in the uppermost 5 m of the canopy. In the wet season, when foliage is most abundant, the horizontal distribution of LAI in a 2100 m² plot ranged widely from 3 to 8, with a mean of 5.41. Changes in mean LAI between wet and dry seasons were small but highly significant. While ca 40% of the area was not affected by local changes in LAI, sizeable small scale changes in LAI did occur between wet and dry season in some locations. Local changes in LAI ranged from –2.3 to 2.4. These changes resulted in a 50% or more increase in light reaching the forest floor at 29% of the measuring locations, and a doubling or more at 13% of the location. Our results imply that structural heterogeneity by simple tree fall gaps do not adequately describe the dynamics of forest canopies.     

鼎湖山顶级森林生态系统水文要素时空规律

运用连续7a(1993～1999)的水文观测资料,对南亚热带顶级生态系统鼎湖山季风常绿阔叶林集水区水文要素时空规律进行分析,得到如下一些主要结论：(1)鼎湖山多年平均降水量为1910mm,湿季降水量占年降水量80％,干季仅占20％。6月份的降水量最大,1月份最小。(2)季风常绿阔叶林冠层截留率为31．8％,湿季的截留量占全年截留量的66．7％,截留量最大值和最小值所在的月份分别为7和1月份。各月的截留率差异很大,截留量大的月份,截留率较低;截留量小的月份,截留率较高。(3)季风常绿阔叶林集水区多年平均总径流量953．0mm,总径流系数49．9％,其中地表径流量为252．3mm,地表径流系数13．2％;地表径流与降水量之间存在二次抛物线型回归关系,与降水强度的关系不大,这说明季风常绿阔叶林的产流形式是是蓄满产流。(4)季风常绿阔叶林多年平均蒸散948．2mm,占同期降水量的49．7％;蒸散力1031．4mm,年蒸散系数为0．92,蒸散月变化规律较降水量的月变化规律有所滞后。(5)系统贮水量的月变化很大,2～8月份,系统处于蓄水阶段;9月份至翌年1月份,系统处于失水阶段。蓄水和失水的最大值分别出现在湿季和干季的第一个月,即4月份和10月份。(6)集水区多年平均水量总输入2129．9mm,实际输入1910mm(降水量),其中219．9mm的水量输入是由系统贮水量变化而产生。支出的总水量2129．9mm,实际支出1901．3mm(径流和蒸散量),其中228．6mm的水量支出是由系统贮水量变化引起的。     

西双版纳热带季节雨林的树种组成和群落结构动态

研究了西双版纳热带季节雨林1 hm2(hectare)动态监测样地1993年与2007年之间树种组成和群落结构的变化。对样地中胸径≥5 cm的乔木进行了每木调查。目前其树种组成的热带分布科、属所占比例分别为91%和94%,具有较高比例的热带植物区系性质。在1993年与2007年两次调查之间,树种数量由145种增至179种,仅有1到2个个体的稀有树种所占比例从54%降为51.1%。从森林的垂直结构来看,A、B、C三层的个体死亡率分别为12.8%、12.9%和19.0%,各层树木的增长率分别为-8.5%、-1.4%和44.8%。与此相对应,C层小径级的树木所占比例有较大提高。虽然小径级的树木在种类和数量上比例增大,但个体数量和种类组成相对稳定的A、B层优势树种变化不大,维持了群落结构的稳定性。14 a间,群落中新增加的具有先锋性质的树种不超过5个。1993年时,A、B两层尚有先锋树种存在,2007年已经从A、B两层中退出。因此,从14 a间树种组成和群落结构的变化来看,虽然具有树木的死亡和增补,但其物种成分和群落结构的总体格局没有明显的变化,处于动态平衡过程中。     

Drivers of productivity and its temporal stability in a tropical tree diversity experiment

There is increasing evidence that mixed‐species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversity–productivity relationships and diversity–stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one‐, two‐, three‐ and five‐species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood‐ and community‐level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability. General patterns were consistent across both scales indicating that tree–tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%–30% in two‐ and three‐species mixtures and 50% in five‐species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño‐related drought. Overall, positive diversity–productivity and diversity–stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed‐species forests may provide both high levels and high stability of production.     

Spatial variability in the vegetation structure and composition of an East African rain forest

We conducted a study of spatial variation in tree community structure and composition in the Kakamega Forest of western Kenya. We compared the tree communities at two sites, Buyangu and Isecheno, separated by approximately 11 km of contiguous forest. All trees ≥15 cm d.b.h. were censused along transects covering 4.95 ha at Buyangu and 4.15 ha at Isecheno. The structure of the forest at the two sites was similar as mean d.b.h. was comparable and stem size class distribution did not differ significantly. However, species richness and stem density were both much higher at Buyangu. The disparity in stem density may be because of the greater abundance at Isecheno of a semi‐woody undergrowth species, Brillantaisia nitens Lindau, believed to inhibit the establishment of tree seedlings. Floristic composition varied strikingly between sites, with 52% of the species occurring only at one site. Densities of those species present at both sites often differed markedly between sites. Potential sources of these intersite differences in floristic composition include small disparities in rainfall, soil composition, elevation, and temperature as well as past differences in anthropogenic disturbance and in large mammal distribution and abundance. Floristic differences between Buyangu and Isecheno appear to be at least partly responsible for the substantial dietary differences between redtail monkeys (Cercopithecus ascanius Audebert) at these sites.     

Spatial and temporal variability of net ecosystem production in a tropical forest: testing the hypothesis of a significant carbon sink

Tropical forest ecosystems play an important role in the global carbon balance. Depending on age and land use, they can act as carbon sources, sinks, or be in approximate balance, but it is uncertain if global environmental changes are forcing these ecosystems outside their natural range of variation. We asked the question of whether or not the net carbon flux of a tropical primary forest, which should be in balance over the long term, is within the expected range of natural variation. A simple Bayesian hypothesis testing method was used to address this question for primary forests in the Porce region of Colombia. Net ecosystem production (NEP) was measured in this forest in a set of 33 permanent plots from 2000 to 2002 in 2, 1‐year intervals. Our estimate of NEP ranged between −4.03 and 2.22 Mg C ha⁻¹ yr⁻¹ for the two intervals. This range was compared with a priori defined range of natural variation estimated from the ecosystem model STANDCARB, which estimated spatial and temporal variation due to gap dynamics. The prior range of variation was estimated between −1.5 and 1.5 Mg C ha⁻¹ yr⁻¹. The observed data on NEP did not provide sufficient evidence to reject the null hypothesis that these forests are in C balance. We concluded that the ecosystem is likely behaving within its range of natural variation, but measurement uncertainties were a major limitation to finding evidence to reject the null hypothesis. A literature review of C flux studies in the tropics revealed that about half of the observations could be explained by gap dynamics alone, while significant C sinks have only been observed during La Niña years, with contrasting results in other tropical forests. In conclusion, observational data of carbon fluxes do not appear to provide direct evidence for a significant carbon sink in some sites in the tropics.     

Palm heart harvesting in the Brazilian Atlantic forest: changes in industry structure and the illegal trade

1. Until the late 1960s, Euterpe edulis was the most important edible palm species in the Brazilian markets. Since then, overharvesting of natural stands and the limited scope of plantations have caused the industry to shift to the multistemmed E. oleracea from the Amazon river estuary.
2. As a result of the relocation of the industry, the structure of the palm harvesting industry has changed and illegal trade has continued to exploit the remaining natural stands of E. edulis .
3. It is suggested that, to avoid the total harvesting of E. edulis and to achieve sustainable management of the palm heart, it will be necessary to diversify the local economy, create incentives for small land owners to improve their management practices, while keeping their competitiveness against the illegal trade, create the incentive of new sources of palm hearts and create a 'green stamping' for management palms.
4. Transformation of the illegal structure of the palm heart industry is probably the only solution to avoid the total depletion of the wild stands of E. edulis .     

Maintenance of agriculture and human habitats within the tropical forest ecosystem

The tactics of shifting cultivation are reviewed with particular reference given to techniques of forest regeneration. The desirability of a forest-fallow system of agriculture is explained. Also discussed are the causes, indicators, and consequences of a breakdown of forest-fallow cultivation. In conclusion, the future of tropical forests for agriculture and habitats for man is considered.This article is a modified version of one originally prepared for a Man and the Biosphere Symposium entitled The Ecological Effects of Increasing Human Activities on Tropical Forest Ecosystems. The Symposium met in Port Moresby, Papua New Guinea, April 28–30, 1975. Proceedings are scheduled for publication in late 1976 and will include another version of the present article.     

三种温带森林大型土壤动物群落结构的时空动态

对帽儿山3种典型森林群落大型土壤动物进行了连续6个月的野外调查研究。通过系统分析,共获得大型土壤动物3604只,隶属于3门6纲17目50科。其中正蚓科(Lumbricidae)、线蚓科(Enchytraeidae)和石蜈蚣目(Lithobiomorpha)为优势类群,常见类群11类。结果表明:(1)水平分布上,密度和生物量红松人工林最高,其次为硬阔叶林,蒙古栎林最少;类群数硬阔叶林最多,蒙古栎林最少;香农指数和丰富度指数均为蒙古栎林最高,红松人工林最低;优势度指数与两者相反;均匀度指数蒙古栎林最高,硬阔叶林最低;(2)垂直分布上,个体密度、类群数及生物量均差异显著(P < 0.001)。3个样地大型土壤动物个体密度表聚性明显;类群数红松人工林自凋落物层向下减少,硬阔叶林和蒙古栎林0-10 cm最多;生物量在0-10 cm土层最大;香农指数随深度增加而减小,优势度指数则相反;(3)在时间变化上,5月和10月个体密度和类群数较多,9月生物量最大;香农指数和优势度指数差异显著(P < 0.01),其他指数各月间无明显差异;(4)与土壤环境因子关系上,总有机碳含量与类群数、个体密度及生物量显著正相关,容重与香农指数显著负相关;典型对应分析结果表明,不同类群大型土壤动物与环境相关性不同。     

Spatiotemporal variability of soil respiration in a seasonal tropical forest

We monitored soil CO ₂ effluxes for over 3 years in a seasonally wet tropical forest in Central Panama using automated and manual measurements from 2013 to 2016. The measurements displayed a high degree of spatial and temporal variability. Temporal variability could be largely explained by surface soil water dynamics over a broad range of temporal scales. Soil moisture was responsible for seasonal cycles, diurnal cycles, intraseasonal variability such as rain‐induced pulses following dry spells, as well as suppression during near saturated conditions, and ultimately, interannual variability. Spatial variability, which remains largely unexplained, revealed an emergent role of forest structure in conjunction with physical drivers such as soil temperature and topography. Mean annual soil CO ₂ effluxes (±SE ) amounted to 1,613 (±59) gC  m^?2 year^?1 with an increasing trend in phase with an El Niño/Southern Oscillation (ENSO ) cycle which culminated with the strong 2015–2016 event. We attribute this trend to a relatively mild wet season during which soil saturated conditions were less persistent.     

Sandfly distribution and abundance in a tropical rain forest

1. The distribution patterns of sandflies (Diptera: Psychodidae) upon tree buttresses were studied in tropical rain forest at Finca la Selva in the Caribbean lowlands of Costa Rica. 2. Four species of sandfly, Lutzomyia shannoni Dyar, L. trapidoi F. & H., L. ylephiletor F. & H. and L. vespertilionis F. & H. comprising 97% of those caught, used tree buttresses as diurnal resting sites. Their distribution on the buttresses was aggregated. 3. During the dry season tree species had no significant effect upon the distribution of the sandflies. However, during the wet season the distributions of two of the species, L. trapidoi and L. ylephiletor, were significantly affected by the species of tree; it is suggested that some species of tree may provide greater protection from rainfall than others. 4. L. vespertilionis was restricted to a single buttress on each positive tree. Distribution of this species is evidently determined by the distribution of its host animal, the bat (Emballonuridae). Female flies feed upon the bat's blood and male flies may be attracted to the bat as it provides a source of female sandflies. It is suggested, therefore, that tree buttresses serve as sandfly swarming sites. 5. Within a large buttress the sandflies are not randomly distributed but are aggregated in particular areas. Within these aggregations, the sandflies are vertically zoned upon the buttress with a shift in species composition with height. Two hypothesis were suggested to account for this distribution pattern: a response to an environmental gradient or an interaction between the four species of fly.     

Site‐to‐site variability and temporal trends of DOC concentrations and fluxes in temperate forest soils

Here, we report site‐to‐site variability and 12–14 year trends of dissolved organic carbon (DOC) from organic layers and mineral soils of 22 forests in Bavaria, Germany. DOC concentrations in the organic layer were negatively correlated with mean annual precipitation and elevation whereas air temperature had a positive effect on DOC concentrations. DOC fluxes in subsoils increased by 3 kg ha^?1 yr^?1 per 100 mm precipitation or per 100 m elevation. The highest DOC concentrations were found under pine stands with mor humus. Average DOC concentrations in organic layer leachates followed the order: pine>oak>spruce>beech. However, the order was different for mean DOC fluxes (spruce>pine>oak>beech) because of varying precipitation regimes among the forest types. In 12 of 22 sites, DOC concentrations of organic layer leachates significantly increased by 0.5 to 3.1 mg C L^?1 yr^?1 during the sampling period. The increase in DOC concentration coincided with decreasing sulfate concentration, indicating that sulfate concentration is an important driver of DOC solubility in the organic layer of these forest sites. In contrast to the organic layer, DOC concentrations below 60 cm mineral soil depth decreased by <0.1–0.4 mg C L^?1 yr^?1 at eight sites. The negative DOC trends were attributed to (i) increasing adsorption of DOC by mineral surfaces resulting from desorption of sulfate and (ii) increasing decay of DOC resulting from decreasing stabilization of DOC by organo‐Al complexes. Trends of DOC fluxes from organic layers were consistent with those of DOC concentrations although trends were only significant at seven sites. DOC fluxes in the subsoil were with few exceptions small and trends were generally not significant. Our results suggest that enhanced mobilization of DOC in forest floors contributed to the increase of DOC in surface waters while mineral horizons did not contribute to increasing DOC export of forest soils.     

海南霸王岭山地原始林与伐后林中木质藤本对支持木的选择

通过对海南霸王岭热带山地原始林与伐后林中树木及其攀附木质藤本的调查,研究原始林与伐后林中木质藤本对支持木的选择性。结果表明:1)6科优势树木中附藤率最高的是野牡丹科(Melastomataceae),附藤率最低的科,原始林中是山矾科(Symplocaceae),伐后林中是茜草科(Rubiaceae)。2)原始林中,谷木(Memecylon ligustrifolium)与线枝蒲桃(Syzygiumaraiocladum)的附藤比率和每木藤本数均高于样地平均水平;三角瓣花(Prismatomeris tetrandra)和龟背灰木(Symplocosandenophylla)的附藤比率均低于样地平均水平,而每木藤本数与样地平均水平之间没有显著差异。伐后林中,谷木的附藤比率和每木藤本数高于样地平均水平;九节(Psychotria rubra)的附藤比率和每木藤本数低于样地平均水平。3)杜仲藤(Parabariummicranthum)的主要支持木是谷木,夜花藤(Hypserpa nitida)的主要支持木是线枝蒲桃。研究表明,木质藤本对支持木在科和种水平上都具有选择性,因此木质藤本会对树木造成不对称影响,进而影响森林的结构和动态。     

Comparing forest structure and biodiversity on private and public land: secondary tropical dry forests in Costa Rica

Secondary forests constitute a substantial proportion of tropical forestlands. These forests occur on both public and private lands and different underlying environmental variables and management regimes may affect post‐abandonment successional processes and resultant forest structure and biodiversity. We examined whether differences in ownership led to differences in forest structure, tree diversity, and tree species composition across a gradient of soil fertility and forest age. We collected soil samples and surveyed all trees in 82 public and 66 private 0.1‐ha forest plots arrayed across forest age and soil gradients in Guanacaste, Costa Rica. We found that soil fertility appeared to drive the spatial structure of public vs. private ownership; public conservation lands appeared to be non‐randomly located on areas of lower soil fertility. On private lands, areas of crops/pasture appeared to be non‐randomly located on higher soil fertility areas while forests occupied areas of lower soil fertility. We found that forest structure and tree species diversity did not differ significantly between public and private ownership. However, public and private forests differed in tree species composition: 11 percent were more prevalent in public forest and 7 percent were more prevalent in private forest. Swietenia macrophylla, Cedrela odorata, and Astronium graveolens were more prevalent in public forests likely because public forests provide stronger protection for these highly prized timber species. Guazuma ulmifolia was the most abundant tree in private forests likely because this species is widely consumed and dispersed by cattle. Furthermore, some compositional differences appear to result from soil fertility differences due to non‐random placement of public and private land holdings with respect to soil fertility. Land ownership creates a distinctive species composition signature that is likely the result of differences in soil fertility and management between the ownership types. Both biophysical and social variables should be considered to advance understanding of tropical secondary forest structure and biodiversity.     

Modelling temporal variability in the carbon balance of a spruce/moss boreal forest

A model of the daily carbon balance of a black spruce/feathermoss boreal forest ecosystem was developed and results compared to preliminary data from the 1994 BOREAS field campaign in northem Manitoba, Canada. The model, driven by daily weather conditions, simulated daily soil climate status (temperature and moisture profiles), spruce photosynthesis and respiration, moss photosynthesis and respiration, and litter decomposition. Model agreement with preliminary field data was good for net ecosystem exchange (NEE), capturing both the asymmetrical seasonality and short-term variability. During the growing season simulated daily NEE ranged from -4 g C m^-2 d^-1 (carbon uptake by ecosystem) to + 2 g C m^-2 d^-1 (carbon flux to atmosphere), with fluctuations from day to day. In the early winter simulated NEE values were + 0.5 g C m^-2 d^-1, dropping to + 0.2 g C m^-2 d^-1 in mid-winter. Simulated soil respiration during the growing season (+ 1 to + 5 g C m^-2 d^-1) was dominated by metabolic respiration of the live moss, with litter decomposition usually contributing less than 30% and live spruce root respiration less than 10% of the total. Both spruce and moss net primary productivity (NPP) rates were higher in early summer than late summer. Simulated annual NEE for 1994 was -51 g C m^-2 y^-1, with 83% going into tree growth and 17% into the soil carbon accumulation. Moss NPP (58 g C m^-2 y^-1) was considered to be litter (i.e. soil carbon input; no net increase in live moss biomass). Ecosystem respiration during the snow-covered season (84 g C m^-2) was 58% of the growing season net carbon uptake. A simulation of the same site for 1968–1989 showed = 10–20% year-to-year variability in heterotrophic respiration (mean of + 113 g C m^-2 y^-1). Moss NPP ranged from 19 to 114 g C m^-2 y^-1; spruce NPP from 81 to 150 g C m^-2 y^-1; spruce growth (NPP minus litterfall) from 34 to 103 g C m^-2 y^-1; NEE ranged from +37 to -142 g C m^-2 y^-1. Values for these carbon balance terms in 1994 were slightly smaller than the 1969–89 means. Higher ecosystem productivity years (more negative NEE) generally had early springs and relatively wet summers; lower productivity years had late springs and relatively dry summers.     

Decelerating growth in tropical forest trees

The impacts of global change on tropical forests remain poorly understood. We examined changes in tree growth rates over the past two decades for all species occurring in large (50-ha) forest dynamics plots in Panama and Malaysia. Stem growth rates declined significantly at both forests regardless of initial size or organizational level (species, community or stand). Decreasing growth rates were widespread, occurring in 24–71% of species at Barro Colorado Island, Panama (BCI) and in 58–95% of species at Pasoh, Malaysia (depending on the sizes of stems included). Changes in growth were not consistently associated with initial growth rate, adult stature, or wood density. Changes in growth were significantly associated with regional climate changes: at both sites growth was negatively correlated with annual mean daily minimum temperatures, and at BCI growth was positively correlated with annual precipitation and number of rainfree days (a measure of relative insolation). While the underlying cause(s) of decelerating growth is still unresolved, these patterns strongly contradict the hypothesized pantropical increase in tree growth rates caused by carbon fertilization. Decelerating tree growth will have important economic and environmental implications.