Post-cultivation Secondary Succession in a Venezuelan Lower Montane Rain Forest

Since tropical rain forests are widely threatened by conversion to agriculture, even within protected areas, an understanding of recovery processes is important for restoration of forest ecosystems and thus conservation of their biodiversity. Secondary succession following land clearance and crop cultivation was studied in a lower montane rain forest in a protected area of the Venezuelan Cordillera de la Costa Central. Forest recovery was studied using a chronosequence of eight 20 × 20 m plots which represented four forest types ca.10 year-old Secondary Forest, ca. 20 year-old Secondary Forest, ca. 35 year-old (uncultivated) secondary forest and mature forest. Species richness and structural complexity increased during succession, with the oldest secondary forest having a physiognomy comparable to the mature forest. Species diversity was lower in the secondary forests than the mature forest, and their floristic composition was distinct. Four phases are hypothesized to occur in the succession process, each with a distinctive species assemblage: initial colonisation by non-woody vegetation; establishment and canopy closure by short-lived small-seeded woody pioneer species; replacement by longer-lived secondary species; and gradual replacement by mature forest large-seeded climax species. Full recovery of the forests in the protected area is likely to take many years, although it may be assisted through conservation management measures.     

Fine Root Distribution in a Lower Montane Rain Forest of Panama

In a Panamanian lower montane rain forest we: (1) analyzed the vertical and horizontal distribution of fine roots; and (2) assessed the relationship of fine root mass to thickness of the soil organic layer, soil pH, and soil-extractable nitrogen. The soil in the study area has developed on volcanic ash deposits and was classified as Hapludand. In randomly distributed samples, the median fine root mass (biomass and necromass, diam ≤ 2 mm) to a depth of 100 cm mineral soil was 544 g/m², 41 percent of which was found in the organic layer. Fine root mass was approximately twice as high in the vicinity of stems of the tree species Oreomunnea mexicana (1069 g/m²) and the palm species Colpothrinax aphanopetala (1169 g/m²) and was associated with thick organic layers. The median thickness of the soil organic layer in a larger random sample ( N = 64) was 8 cm with a considerable variation (interquartile range: 7 cm). In these samples, the density of fine root biomass was correlated with the concentration of extractable nitrogen ( r = 0.33, P = 0.011), and on an areal basis, fine root biomass in the organic layer increased with increasing thickness of the organic layer ( r = 0.63, P < 0.001) and decreasing pH_KCl ( r =−0.33, P < 0.01). Fine root biomass in the upper mineral soil did not show significant correlations with any of the studied parameters.     

西双版纳热带山地雨林群落学特征分析

根据4块样地的调查资料,分析了西双版纳热带山地雨林的群落学特征。结果表明：在2500m^2的样地中,共计有维管束植物105～178种,分属于46～61科、80～130属。区系成分可分为11个类型,4块样地均以热带区系成分为主（约90％）,其中又以热带亚洲（印度-马来西亚）分布（23．40％～32．88％）和泛热带分布（25．20％～36．67％）的成分占优势。群落以高位芽植物为主,其中又以中（约50％）、小高位芽植物（21．30％～27．78％）占优势;在叶片特征上,群落以中叶（72．66％～82．54％）、单叶（75％～86．96％）、纸质（58．73％～62．22％）、全缘（76．67％～83．33％）、非尾尖（约90％）为主。低海拔带上山地雨林的热带区系成分以及藤本植物、中叶和全缘叶植物所占的比例高于高海拔带上的山地雨林,而小高位芽植物比例则低于后者,体现出较强的热带雨林性质。西双版纳的山地雨林与本地区的沟谷雨林、低丘雨林相比表现为热带区系成分、藤本植物、大高位芽植物以及巨叶、大叶植物较少,而温带成分、小高位芽植物、小叶植物较为丰富,体现出向常绿阔叶林的过渡特征。     

The Hummingbird Plant Community of a Tropical Montane Rain Forest in Southern Ecuador

Abstract: The diversity of a hummingbird plant community in the eastern Andes of southern Ecuador was studied on the equivalent of a hectare (two 500 ' 10 m transects) at 1920 - 2100 m a.s.l. over the course of a year. A total of 3186 flowering individuals, representing 12 plant families, 29 genera and 72 species, were found to be visited by hummingbirds. Bromeliaceae had the most species visited, followed by Orchidaceae and Ericaceae. The majority of visited plant species were represented by a very few individuals, and only a few species of the Bromeliaceae and Orchidaceae appeared in large numbers of visited individuals. With regard to life forms visited by hummingbirds, epiphytes predominated (59 %), followed by trees and shrubs (29 %), vines (8 %) and herbs (4 %). Visited flowers usually had short- to medium-long floral tubes which were either functionally or morphologically tubiform or campanulate. Fifty percent of the species had red-coloured flowers, and a considerable number of the blossoms (43 %) displayed contrasting colours. The 72 plant species received visits from 26 species of Trochilidae (hummingbirds) and two species of Coerebidae (honeycreepers). A mere eight species of hummingbirds were seen frequently at the study area; the remaining species were only occasionally sighted. The eight frequently sighted species of hummingbirds made use of a total of 74 % of all hummingbird-visited plant species growing in the study area.     

Tree Diversity,Forest Structure and Productivity along Altitudinal and Topographical Gradients in a Species-Rich Ecuadorian Montane Rain Forest

We studied the spatial heterogeneity of tree diversity, and of forest structure and productivity in a highly diverse tropical mountain area in southern Ecuador with the aim of understanding the causes of the large variation in these parameters. Two major environmental gradients, elevation and topography, representing a broad range of climatic and edaphic site conditions, were analyzed. We found the highest species richness of trees in valleys <2100 m. Valleys showed highest values of basal area, leaf area index and tree basal area increment as well. Tree diversity also increased from ridges to valleys, while canopy openness decreased. Significant relationships existed between tree diversity and soil parameters (pH, total contents of Mg, K, Ca, N and P), and between diversity and the spatial variability of pH and Ca and Mg contents suggesting a dependence of tree diversity on both absolute levels and on the small-scale heterogeneity of soil nutrient availability. Tree diversity and basal area increment were positively correlated, partly because both are similarly affected by soil conditions. We conclude that the extraordinarily high tree species richness in the area is primarily caused by three factors: (1) the existence of steep altitudinal and topographic gradients in a rather limited area creating a small-scale mosaic of edaphically different habitats; (2) the intermingling of Amazonian lowland plant species, that reach their upper distribution limits, and of montane forest species; and (3) the geographical position of the study area between the humid eastern Andean slope and the dry interandean forests of South Ecuador.     

Reproductive biology of two tree species of Leguminosae in a Montane Rain Forest in southeastern Brazil

Studies on sympatric species that belong to the same taxonomic group and have the same pattern of pollination and seed dispersion, but different patterns of geographical distribution and structural parameters may be worthwhile to understand the evolution of their compatibility system. We studied the floral biology, pollinators and reproductive systems of two tree species in Itatiaia, an area covered with montane rain forest. Pseudopiptadenia contorta is widespread in Brazil but is present at a low relative density at the study site, while Pseudopiptadenia leptostachya is restrictively distributed but presents one of its highest relative density values in Itatiaia. Both species have small flowers arranged in compact spicate raceme inflorescences that constitute the pollination unit. Pollinators are native stingless bees and European honeybees that collect pollen from their nectarless flowers. Although both species have similar floral characteristics and pollination system, their compatibility system differs. Fruit set after manual pollinations and observation of the pollen tube growth showed that P. leptostachya is self-compatible while P. contorta only set fruits after cross-pollinations, and is thus a self-incompatible species. However, fruit set did not differ between the species, indicating that structural parameters are not affected by the compatibility system. Thus, the difference between these two species in terms of reproductive system may be related to other factors such as patterns of geographical distribution.     

A Technique to Identify Changes in Hemlock Forest Health over Space and Time Using Satellite Image Data

The objective of this study was to develop a technique to classify health of eastern hemlock stands using historical satellite images. While remote sensing and geographic information systems have been used successfully to classify forest health using recent images, applying this process to older images is problematic because contemporaneous field data are not available to measure the accuracy of the classification of historical images. Data ranges were established for each hemlock health class using a contemporary image and field data. These ranges were used to level-slice archived images to create a series of health-class maps that show changes in forest health over time. By applying cross-tabulation procedures to pairs of classified images, it is possible to construct a transition map that indicates how the hemlock health class of each pixel in the images of the study area has changed over time. The resulting maps provide a look back at forest conditions of the past and can be used to identify areas of special interest.     

Water Relations and CO2 Exchange of Tropical Bryophytes in a Lower Montane Rain Forest in Panama

For 6 tropical bryophytes, measurements of the diel courses in water status and net CO₂ exchange were made in a submontane tropical rain forest in Panama. In addition, the response of gas exchange to changes in photon flux density (PPFD) and thallus water content (WC) was studied under controlled conditions. Diel variation of WC was pronounced, and both low and high WC limited carbon gain considerably. Low PPFD, e.g. during rain storms, was less important in limiting CO₂ exchange. More than half of the mean diurnal carbon gain of 2.9 mg C per g thallus carbon was lost during the night as respiration. Assuming that the average 24-h carbon gain was representative for the entire year, we estimated the net annual primary productivity of the mosses and liverworts to be 45% of the initial plant carbon content.     

海南霸王岭热带山地雨林森林循环与树种多样性动态

通过对海南岛霸王岭热带山地雨林的调查 ,研究了热带山地雨林树种多样性特征随森林循环的动态变化规律。结果表明 :( 1 )热带山地雨林森林循环不同阶段斑块在森林景观中所占的面积比例分别是 :林隙阶段 ( G)占 38.5 0 % ,建立阶段 ( B)占 2 8.5 0 % ,成熟阶段 ( M)占 2 7.0 0 % ,衰退阶段 ( D)占 6 .0 0 %。 ( 2 )热带山地雨林中乔木树种的密度随森林循环的变化趋势是由 G→B→M呈现出逐渐增加的趋势 ,以成熟阶段达到最大 ,而到衰退阶段又趋于下降。灌木树种则表现出 G阶段斑块的密度最大 ,B阶段的最小 ,从 B到 M有所增加 ,到 D又稍有下降。 ( 3)热带山地雨林中不同高度级和不同径级的树木的密度在森林循环的不同阶段表现出不同的增减趋势 ,其随森林循环过程呈现出的动态变化可能与不同阶段斑块内的空间、环境及物种生物学特性有关。 ( 4 )热带山地雨林中树木的平均胸径、平均高、平均胸高断面积、平均单株材积随森林循环过程呈现出不断增加的趋势 ,其中平均胸径和平均高随森林循环的变化较为平缓 ,而平均胸高断面积和平均单株材积之变化较为陡急。 ( 5 )热带山地雨林森林循环不同阶段的物种多样性指数不同 ,其中 G和 B阶段的物种丰富度和多样性指数值较接近 ,M阶段的物种丰富度达到最大 ,D阶段则最小。     

Mycorrhizal,Endophytic and Ecomorphological Status of Tree Roots in the Canopy of a Montane Rain Forest

Our investigation provides data on the symbiotic status and the functional morphology of tree fine roots in canopy soils of an upper montane forest in Costa Rica. Canopy roots completely lacked mycorrhizal colonization, but showed a high abundance of endophytic fungi suggesting a reduced ability for nutrient acquisition compared with terrestrial roots.     

Asymbiotic Nitrogen Fixation and Litter Decomposition on a Long Soil-Age Gradient in Hawaiian Montane Rain Forest1

We determined rates of decomposition and asymbiotic nitrogen fixation in the leaf litter of Cheirodendron spp. on the Hawaiian Islands. Leaf litter was collected from four sites on a long soil-age gradient (300 yr to 4.1 M yr) and decomposed at two sites that differed widely in substrate age and nutrient availability. Rates of decomposition were higher in litter decomposed at the older site, where nutrient availability was greater. A substantial amount of nitrogen and phosphorus immobilization occurred in litter decomposed at the older site, with more immobilization occurring in litter with lower initial nitrogen and phosphorus concentrations, suggesting both supply and demand controls on nutrient immobilization. Potential rates of nitrogen fixation were very low in the first 25 d (0–5 nmol acetylene/gdw/h), rose to much higher rates by 70 d (20–45 nmol), and then declined by 140 d. We found no significant difference in rates of potential nitrogen fixation between sites of decomposition, but there was a strong substrate effect, with higher rates in litter with low lignin, low nitrogen, and high phosphorus. Where significant immobilization of nitrogen occurred for decomposing Cheirodendron, nitrogen fixation could have comprised no more than 10 percent of immobilized nitrogen. Overall, rates of nitrogen fixation were dependent on the source of the decomposing substrate but not on the site of decomposition, while short-term decomposition and nutrient immobilization were strongly dependent on the site of decomposition but not as much on the source of the decomposing substrate.     

Diets of Frugivorous Bats in Montane Rain Forest and Coffee Plantations in Southeastern Chiapas,Mexico

Frugivorous are one of the main diaspore dispersers in tropical ecosystems, particularly in open areas and sites in the early stages of ecological succession. Frugivorous bat species respond differentially to habitat modification, and in the context of their diaspore dispersal functions it is important to understand species' ecological requirements. We compared the diversity of diaspores, obtained from fecal samples and from fruits carried by frugivorous bats, among five shaded coffee plantations under different management regimes and a montane rain forest in southeastern Chiapas, Mexico. At each site, bats were captured every 2 mo from March 2004 to July 2005, using six mist‐nets, during two consecutive nights. We captured 2589 individuals from 18 frugivorous species, from which we collected 969 fecal samples, containing 42 diaspore species associated with early and late successional plants. Although, we captured more frugivorous bat species in montane rain forest, the number of diaspore species in this site (N=14) was not significantly different from the coffee plantations under different management regimes (16–24). In montane rain forest, Sturnira ludovici fed mainly on Piper auritum, but in coffee plantations ate Peperomia sp., Saurauia madrensis, Solanum chrysotrichum and Solanum diphyllum. Artibeus jamaicensis and Artibeus intermedius feed mostly Cecropia obtusifolia and Ficus cookii in all coffee plantations. We suggest that the presence of frugivorous bats in shaded coffee plantations is favored by trees and shrubs associated with secondary and introduced vegetation that farmers have allowed to grow within or around the plantations.     

Gap disturbance regime and composition in the Atlantic Montane Rain Forest: the influence of topography

In the Atlantic Montane Rain Forest of south-eastern Brazil, a field study was carried out to describe the forest disturbance regime, analyse canopy gap composition and evaluate the influence of habitat parameters on gap tree species composition. We characterized canopy gaps considering the group of variables as follows: area, type and number of tree/branch falls, topographic position, soil coverage and surrounding canopy trees. Gap composition was assessed at species level by measuring all individuals inside gaps higher than one meter. Mean gap area of the 42 canopy gaps analysed was 71.9 ± 9.0 m2 (mean ± SE). Out of the studied gaps, 35.7% were created by uprooted and by snapped trees, 16.7% by dead-standing trees and 11.9% by the fall of large branches. The disturbance regime was characterized by gap openings predominantly smaller than 150 m2 and by spatial patterning related to topography. Ridges had smaller gaps and higher proportions of gaps created by branch falls; slopes had bigger gaps generally created by uprooting events. The more abundant and frequent species were shade tolerant and the more species-rich families found inside gaps did not differ from the forest as a whole. Pioneer species were rare and restricted to medium and large size classes. The Indicator Species Analysis and the Canonical Correspondence Analysis indicated gap area, topography and the percentage of soil cover by the genera Calathea and Ctenanthe were the predominant variables correlated with woody species distribution. So, topography emerged as an important issue not only to the gap disturbance regime, but also to gap colonization. In respect to the influence of gap processes on the Atlantic Montane Rain Forest regeneration, our results support the view that canopy gap events may not be working as promoters of community wide floristic shifts.     

Nitrogen Fixation in Bryophytes, Lichens, and Decaying Wood along a Soil-age Gradient in Hawaiian Montane Rain Forest

We determined rates of acetylene reduction and estimated total nitrogen fixation associated with bryophytes, lichens, and decaying wood in Hawaiian montane rain forest sites with underlying substrate ranging in age from 300 to 4.1 million years. Potential N fixation ranged from ca 0.2 kg/ha annually in the 300‐year‐old site to ca 1 kg/ha annually in the 150,000‐year‐old site. Rates of acetylene reduction were surprisingly uniform along the soil‐age gradient, except for high rates in symbiotic/associative fixers at the 150,000‐year‐old site and in heterotrophic fixers at the 2100‐year‐old site. Low fixation at the youngest site, where plant production is known to be N‐limited, suggests that demand for N alone does not govern N fixation. Total N fixation was highest in sites with low N:P ratios in leaves and stem wood, perhaps because epiphytic bryophytes and lichens depend on canopy leachate for mineral nutrients and because heterotrophic fixation is partly controlled by nutrient supply in the decomposing substrate; however, differences in substrate cover, rather than in fixation rates, had the largest effect on the total N input from fixation at these sites.     

Soil Phosphorus Fractionation and Phosphorus-Use Efficiency of a Bornean Tropical Montane Rain Forest During Soil Aging With Podozolization

We compared phosphorus (P) dynamics and plant productivity in two montane tropical rain forests (Mount Kinabalu, Borneo) that derived from similar parent materials (largely sedimentary rocks) and had similar climates but differed in terms of soil age. The younger site originated from Quaternary colluvial deposits, whereas the older site had Tertiary-age material. The older site had a distinctive spodic horizon, reduced levels of labile inorganic soil P, higher concentrations of recalcitrant organic soil P, and lower rates of net soil N mineralization. P fertilization led to soil nitrogen (N) immobilization in the P-deficient soil, indicating that soil N mineralization was limited by P at the P-deficient older site. Mean foliar nutrient concentration (on both a weight and an area basis) was similar at the two sites for all elements except P, which was lower at the older site. Aboveground net primary production (ANPP) was lower at the older site than at the younger one; this difference could be explained by the reduced availability of P and N (as down-regulated by P) at the older site. The relatively ample allocation of P and N to leaves, despite the reduced availability at the P-deficient old site, was attributable to its high resorption efficiency. High resorption resulted in lower concentrations of elements in leaf litter—that is, less decomposable low-quality litter. On the other hand, the concentration of leaf litter lignin was considerably lower at the older site; this appeared to be a de facto adaptive mechanism to avoid retarding litter decomposition.     

海南岛热带山地雨林演替后期幼树的叶片特征

植物的生理生态特征决定了植物在生态系统中的分布模式和物种的丰度。本文在开展样地调查的基础上,应用Li-6400光合测定系统研究了海南岛热带山地雨林原始林3个层次的6个优势种与9个伴生种幼树的光合作用参数,并测定了相应叶片的比叶面积（SLA）和氮、磷含量。15个种的SLA为38.2~143.7 cm²·g^-1、单位面积最大光合速率（A_-area）为2.77~7.61 μmol·m^-2·s^-1、单位干重最大光合速率（A_-mass）为21.2~83.4 μmol·kg^-1·s^-1,单位面积暗呼吸速率（R_d-area）为0.02~1.15 μmol·m^-2·s^-1、单位干重暗呼吸速率（R_d-mass）为0.19~12.61 μmol·kg^-1·s^-1,光补偿点（LCP）为2.5~32.2 μmol·m^-2·s^-1,表观量子效率（Ф）为0.006 6~0.042 8 μmol·μmol^-1 photon,叶片氮（LN）含量为7.98~23.63 mg·g^-1,叶片磷（LP）含量为3.98~13.40 mg·g^-1。中上层种的幼树具有较高的比叶面积、单位干重最大光合速率和表观量子效率;上层种的暗呼吸速率最低;幼树叶片的氮、磷含量随物种成树所在层次升高而减少,次序为下层种>中层种>上层种,但差异均不显著（P>0.05）。优势种和伴生种的光响应参数与SLA存在相关关系。伴生种的LN与SLA正相关（P<0.01）,LP与SLA负相关（P<0.05）;优势种的LN、LP与SLA不相关（P>0.05）。     

西双版纳勐宋热带山地雨林种子植物区系的初步研究

西双版纳勐宋的热带山地雨林是一种东南亚热带山地雨林的北缘类型,它在植物区系组成上和群落学特征上是介于热带雨林和南亚热带常绿阔叶林之间的热带山地常绿阔叶林.该区系有种子植物116科327属623种.科、属、种的分布区类型分析表明:科的分布区类型以热带和亚热带分布最多(32.8%),其次是热带到温带分布(23.3%)和典型热带分布(18.1%);属的分布区类型以热带亚洲分布最多(28.4%),其次是泛热带分布(26.8%),热带分布多于温带分布(258:55);种的分布区类型以热带亚洲分布最多(65.0%),其次是中国特有分布(26.5%),热带分布远多于温带分布(431:18).区系的热带性质显著,具有明显的印度-马来西亚植物区系特点,属于热带亚洲区系的一部分.由于特殊的地理位置,西双版纳地区是许多典型热带植物的分布北界,同时又是几种热带成分的交汇地带.     

Environmental and Biotic Controls over Aboveground Biomass Throughout a Tropical Rain Forest

The environmental and biotic factors affecting spatial variation in canopy three-dimensional (3-D) structure and aboveground tree biomass (AGB) are poorly understood in tropical rain forests. We combined field measurements and airborne light detection and ranging (lidar) to quantify 3-D structure and AGB across a 5,016 ha rain forest reserve on the northeastern flank of Mauna Kea volcano, Hawaii Island. We compared AGB among native stands dominated by Metrosideros polymorpha found along a 600–1800 m elevation/climate gradient, and on three substrate-age classes of 5, 20, and 65 kyr. We also analyzed how alien tree invasion, canopy species dominance and topographic relief influence AGB levels. Canopy vertical profiles derived from lidar measurements were strong predictors (r ² = 0.78) of AGB across sites and species. Mean AGB ranged from 48 to 363 Mg ha⁻¹ in native forest stands. Increasing elevation corresponded to a 53–84% decrease in AGB levels, depending upon substrate age. Holding climate constant, changes in substrate age from 5 to 65 kyr corresponded to a 23–53% decline in biomass. Invasion by Psidium cattleianum and Ficus rubiginosa trees resulted in a 19–38% decrease in AGB, with these carbon losses mediated by substrate age. In contrast, the spread of former plantation tree species Fraxinus uhdei corresponded to a 7- to 10-fold increase in biomass. The effects of topographic relief at both local and regional scales were evident in the AGB maps, with poorly drained terrain harboring 76% lower biomass than forests on well-drained relief. Our results quantify the absolute and relative importance of environmental factors controlling spatial variation in tree biomass across a rain forest landscape, and highlight the rapid changes in carbon storage incurred following biological invasion. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  GPA and RFH conceived of or designed the study. GPA, RFH, TAV, DEK, and TKB performed research and analyzed data. GPA, RFH, DEK, and TKB contributed new methods or models. GPA wrote the article.     

Tropical Rain Forest Recovery from Cyclone Damage and Fire in Samoa1

In 1990 and 1991, Samoa was struck by two cyclones, Ofa and Val. In the Tafua Rain Forest Preserve on the island of Savai'i, one part of the forest also burned after the first cyclone. Here we report on patterns of regeneration and changes in tree species composition in the Tafua lowland rain forest after five years of recovery from cyclone and fire disturbance. In the unburned area, tree canopy cover increased from 27 percent after the last cyclone to 58 percent, and in the burned area from below 12 to 49 percent. Nine of the ten most common tree species decreased in relative abundance in the entire forest after the last cyclone. One fast growing pioneer species, Macaranga harveyana now makes up 42 percent of the total number of trees (>5 cm DBH) in the unburned area and 86 percent in the burned area. Large interspecific differences occur in size distribution and there are at least four distinguishable regeneration patterns, which may be related to shade tolerance. Mean number of species per plot was generally higher in the unburned area than in the burned area, while the Shannon evenness index was higher in the unburned than in the burned area only for trees above 1 cm DBH. Species with fruits known to be fed upon by birds and/or bats generally made up a larger proportion of all trees in the burned than in the unburned area. In contrast to other studies of post‐cyclone regeneration, in which recovery is often rapid due to resprouting of trees, recovery in the Tafua forest was a slow process with regeneration more dependent on vertebrate seed dispersal than on resprouting.     

The Quantity and Turnover of Dead Wood in Permanent Forest Plots in Six Life Zones of Venezuela1

Dead wood can be an important component of the carbon pool in many forests, but few measurements have been made of this pool in tropical forests, To fill this gap, we determined the quantity of dead wood (downed and standing dead) in 25 long-term (up to 30 yr) permanent forest plots located in six different life zones of Venezuela. Downed wood was separated into fine (< 10 cm in diameter) and coarse (≥ 10 cm in diameter) classes, and three decomposition states (sound, intermediate, or rotten). The total quantity of dead wood, averaged by life zone, was lowest in the dry (2.43 Mg/ha), reached a peak in the moist (42.33 Mg/ha) and decreased slightly in the wet (34.50 Mg/ha) life zone. Most of the dead wood was in the standing dead category (about 42–76% of the total). The decomposition state of dead wood in all plots was mostly rotten (45%) or intermediate (44%); there was little sound wood (11%). Turnover rates of dead wood generally ranged between 0.03/yr to 0.52/yr with no clear trend with life zone. The large amount of dead wood in some plots was equivalent to about 20 percent or less of aboveground biomass, indicating that dead wood can represent a significant amount of carbon in these forests.