Transport and particle size distribution of suspended and benthic organic matter in three headwater streams

The breakdown and decomposition of two species of deciduous leaf litter, Fagus sylvatica L. and Salix viminalis L. and two species of aquatic macrophyte Isoetes lacustris L. and Potamogeton perfoliatus L. were examined in an oligotrophic lake. In all cases plant litter in coarse mesh litter bags lost significantly more material than the fine mesh after 1 years submergence in the lake. This however was considered to be the result of physical environmental factors and microbial activity rather than animal processing. The litter was ranked in order of fastest to slowest rates of decay as follows — Isoetes, Potamogeton, Salix and Fagus. Decomposition processes proceeded at a relatively slow rate as a result of low temperatures and low phosphate and mineral ion concentration. The results suggested that there was an accumulation of organic material in the lake.     

Dry matter and nutrient inputs through litter fall in a dry tropical forest of India

The present paper elucidates the pattern of leaf and non-leaf fall and quantifies of the total annual input of litter in a dry tropical forest of India. In addition, concentration of selected nutrients in various litter species and their annual return to the forest floor are examined. Total annual input of litter measured in litter traps ranged between 488.0–671.0 g m^-2 of which 65–72% was leaf litter fall and 28–35% wood litter fall. 73–81% leaves fall during the winter season. Herbaceous litter fall ranged between 80.0–110.0 g m^-2 yr^-1. The annual nutrient return through litter fall amounted (kg ha^-1): 51.6–69.6 N, 3.1–4.3 P, 31.0–40.0 Ca, 14.0–19.0 K and 3.7–5.0 Na, of which 71–77% and 23–29% were contributed by leaf and wood litter fall, respectively for different nutrients. Input of nutrients through herbaceous litter was: 13.0–16.6 for N, 1.0–1.4 for P, 4.0–5.0 for Ca, 7.9–10.5 for K and 0.8–1.0 kg ha^-1 yr^-1 for Na.     

亚高山森林溪流镉储量与分配的动态变化特征

镉(Cd)是一种有害重金属元素,能够伴随溪流水体流动和物质沉积影响下游流域的生态环境安全,但缺乏必要关注。为了解森林溪流Cd储量及其分配的动态变化特征,以岷江上游亚高山森林集水区的溪流为研究对象,在长度10—50、50—150、150—260m区间内各选取5条典型溪流,研究Cd元素在亚高山森林-溪流-河流集合生态系统中的迁移过程。结果表明:亚高山森林溪流的Cd储量介于2.57—128.46mg/m²之间,主要储存于沉积物中;森林溪流上、中、下游的Cd储量没有显著差异;森林溪流的Cd储量以秋季凋落物高峰期最高,春季凋落物高峰期最低;森林溪流的上、中、下游Cd储量均在秋季凋落高峰最高,上、中游在春季凋落高峰最低,下游在非凋落高峰最低;凋落物的Cd储量与溪流水文特征密切相关。可见,亚高山森林溪流Cd储量动态具有季节性变化和一定的自净能力,这些结果为进一步了解Cd元素在水-陆生态系统的生物地球化学循环提供了新的角度。     

Altered litter inputs modify carbon and nitrogen storage in soil organic matter in a lowland tropical forest

Biogeochemistry - Soil organic matter (SOM) in tropical forests is an important store of carbon (C) and nutrients. Although SOM storage could be affected by global changes via altered plant...     

Iron oxidation stimulates organic matter decomposition in humid tropical forest soils

Humid tropical forests have the fastest rates of organic matter decomposition globally, which often coincide with fluctuating oxygen (O₂) availability in surface soils. Microbial iron (Fe) reduction generates reduced iron [Fe(II)] under anaerobic conditions, which oxidizes to Fe(III) under subsequent aerobic conditions. We demonstrate that Fe (II) oxidation stimulates organic matter decomposition via two mechanisms: (i) organic matter oxidation, likely driven by reactive oxygen species; and (ii) increased dissolved organic carbon (DOC) availability, likely driven by acidification. Phenol oxidative activity increased linearly with Fe(II) concentrations (P < 0.0001, pseudo R² = 0.79) in soils sampled within and among five tropical forest sites. A similar pattern occurred in the absence of soil, suggesting an abiotic driver of this reaction. No phenol oxidative activity occurred in soils under anaerobic conditions, implying the importance of oxidants such as O₂ or hydrogen peroxide (H₂O₂) in addition to Fe(II). Reactions between Fe(II) and H₂O₂ generate hydroxyl radical, a strong nonselective oxidant of organic compounds. We found increasing consumption of H₂O₂ as soil Fe(II) concentrations increased, suggesting that reactive oxygen species produced by Fe(II) oxidation explained variation in phenol oxidative activity among samples. Amending soils with Fe(II) at field concentrations stimulated short‐term C mineralization by up to 270%, likely via a second mechanism. Oxidation of Fe(II) drove a decrease in pH and a monotonic increase in DOC; a decline of two pH units doubled DOC, likely stimulating microbial respiration. We obtained similar results by manipulating soil acidity independently of Fe(II), implying that Fe(II) oxidation affected C substrate availability via pH fluctuations, in addition to producing reactive oxygen species. Iron oxidation coupled to organic matter decomposition contributes to rapid rates of C cycling across humid tropical forests in spite of periodic O₂ limitation, and may help explain the rapid turnover of complex C molecules in these soils.     

海南岛霸王岭热带山地常绿林和热带山顶矮林群落特征

热带山地常绿林和热带山顶矮林均属于热带云雾林.为了揭示其群落结构和物种多样性特征,在海南岛霸王岭热带山地常绿林和热带山顶矮林分别设置8个和10个2,500 m2样方,调查所有DBH≥1cm的乔木、灌木和藤本植株.结果显示:(1)热带山地常绿林幼树(1cm≤DBH＜5 cm)和小树(5cm≤DBH＜ 10 cm)的平均密...     

Above- and belowground organic matter storage and production in a tropical pine plantation and a paired broadleaf secondary forest

The distribution of tree biomass and the allocation of organic matter production were measured in an 11-yr-old Pinus caribaea plantation and a paired broadleaf secondary forest growing under the same climatic conditions. The pine plantation had significantly more mass aboveground than the secondary forest (94.9 vs 35.6 t ha^-1 for biomass and 10.5 vs 5.0 t ha^{-1 for litter}), whereas the secondary forest had significantly more fine roots (⩽2 mm diameter) than the pine plantation (10.5 and 1.0 t ha^-1, respectively). Standing stock of dead fine roots was higher than aboveground litter in the secondary forest. In contrast, aboveground litter in pine was more than ten times higher than the dead root fraction. Both pine and secondary forests had similar total organic matter productions (19.2 and 19.4 t ha^-1 yr^-1, respectively) but structural allocation of that production was significantly different between the two forests; 44% of total production was allocated belowground in the secondary forest, whereas 94% was allocated aboveground in pine. The growth strategies represented by fast growth and large structural allocation aboveground, as for pine, and almost half the production allocated belowground, as for the secondary forest, illustrate equally successful, but contrasting growth strategies under the same climate, regardless of soil characteristics. The patterns of accumulation of organic matter in the soil profile indicated contrasting nutrient immobilization and mineralization sites and sources for soil organic matter formation.     

Influence of hydrological pathways on dissolved organic carbon fluxes in tropical streams

Water flow pathways and water balance are fundamental components for understanding the dynamics of C in the soil/water interface of small basins. The objective of this study was to describe the seasonal variations and estimate the annual balance of dissolved organic carbon (DOC ) by comparing two tropical microbasins (preserved forest—PF and cacao plantation—CP ). Twenty‐one weekly collections were conducted from September to December 2012 and from April to June 2013. The calculation of the partial balance considered precipitation (P) as inflow and the stream as outflow. The samples were filtered and analyzed using a TOC analyzer. Overall, the DOC was higher CP compared with FP . The behavior of both venues showed that rainy season caused an increase in concentrations in the overland flow (OF ) and in the stream, and a decrease in the precipitation (P) and in the throughfall (T). In the CP , the outflow and the soil were chiefly responsible for the high DOC concentrations in the stream, when compared to the PF , which is the result of constant OM decomposition. Soil composition contributes to the control of DOC consumption in each type of soil. The balances were negative in both microbasins, although losses were higher in the AFS (agroforestry systems) when compared to the PF , especially during rainy seasons (?8.98 and ?3.05 kg ha^?1 year^?1, CP and FP , respectively). Thus, the high annual loss of DOC in the CP of the microbasins during the rainy season indicates changes in ecosystem metabolism due to the vegetation cover and to the interactions with the soil.     

基于FORCCHN模型的中国典型森林生态系统碳通量模拟

森林生态系统是陆地碳循环的重要组成部分,其固碳能力显著高于其他陆地生态系统,研究森林生态系统碳通量是认识和理解全球变化对碳循环影响的关键。碳循环模型是研究森林生态系统碳通量有效工具。以长白山温带落叶阔叶林、千烟洲亚热带常绿针叶林、鼎湖山亚热带常绿阔叶林和西双版纳热带雨林等4种中国典型森林生态系统为研究对象,利用涡度相关2003-2012年观测数据,评估FORCCHN模型对生态系统呼吸（ER）,总初级生产力（GPP）,净生态系统生产力（NEP）的模型效果。结果表明：（1） FORCCHN模型能够较好的模拟中国4种典型森林生态系统不同时间尺度的碳通量。落叶阔叶林和常绿针叶林ER和GPP的逐日变化模拟效果较好（ER的相关系数分别为0.94和0.92,GPP的相关系数分别为0.86和0.74）;（2）4种森林生态系统碳通量季节动态模拟值和观测值显著相关（P<0.01）,ER、GPP、NEP的观测值和模拟值的R²分别为0.77-0.93、0.54-0.88和0.15-0.38;模型可以很好地模拟森林生态系统不同季节碳汇（NEP>0）,碳源（NEP<0）的变化规律;（3）4种森林生态系统碳通量模拟值与观测值的年际变化有很好的吻合度,但在数值大小上存在差异,模型高估了常绿阔叶林的ER和GPP,略微低估了其他3种森林生态系统ER和GPP。     

Effects of logging on carbon dynamics of a jack pine forest in Saskatchewan,Canada

We calculated carbon budgets for a chronosequence of harvested jack pine (Pinus banksiana Lamb.) stands (0‐, 5‐, 10‐, and～29‐year‐old) and a～79‐year‐old stand that originated after wildfire. We measured total ecosystem C content (TEC), above‐, and belowground net primary productivity (NPP) for each stand. All values are reported in order for the 0‐, 5‐, 10‐, 29‐, and 79‐year‐old stands, respectively, for May 1999 through April 2000. Total annual NPP (NPP_T) for the stands (Mg C ha^?1 yr^?1±1 SD) was 0.9±0.3, 1.3±0.1, 2.7±0.6, 3.5±0.3, and 1.7±0.4. We correlated periodic soil surface CO₂ fluxes (R_S) with soil temperature to model annual R_S for the stands (Mg C ha^?1 yr^?1±1 SD) as 4.4±0.1, 2.4±0.0, 3.3±0.1, 5.7±0.3, and 3.2±0.2. We estimated net ecosystem productivity (NEP) as NPP_T minus R_H (where R_H was calculated using a Monte Carlo approach as coarse woody debris respiration plus 30–70% of total annual R_S). Excluding C losses during wood processing, NEP (Mg C ha^?1 yr^?1±1 SD) for the stands was estimated to be ?1.9±0.7, ?0.4±0.6, 0.4±0.9, 0.4±1.0, and ?0.2±0.7 (negative values indicate net sources to the atmosphere.) We also calculated NEP values from the changes in TEC among stands. Only the 0‐year‐old stand showed significantly different NEP between the two methods, suggesting a possible mismatch for the chronosequence. The spatial and methodological uncertainties allow us to say little for certain except that the stand becomes a source of C to the atmosphere following logging.     

Allochthonous litter inputs, organic matter standing stocks, and organic seston dynamics in upland Panamanian streams: potential effects of larval amphibians on organic matter dynamics

Allochthonous inputs of detritus represent an important energy source for streams in forested regions, but dynamics of these materials are not well studied in neotropical headwater streams. As part of the tropical amphibian declines in streams (TADS) project, we quantified benthic organic matter standing stocks and organic seston dynamics in four Panamanian headwater streams, two with (pre-amphibian decline) and two without (post-decline) healthy amphibian assemblages. We also measured direct litterfall and lateral litter inputs in two of these streams. Continuous litterfall and monthly benthic samples were collected for 1 year, and seston was collected 1–3 times/month for 1 year at or near baseflow. Direct litterfall was similar between the two streams examined, ranging from 934–1,137 g DM m⁻² y⁻¹. Lateral inputs were lower, ranging from 140–187 g DM m⁻¹ y⁻¹. Dead leaves (57–60%), wood (24–29%), and green leaves (8–9%) contributed most to inputs, and total inputs were generally higher during the rainy season. Annual habitat-weighted benthic organic matter standing stocks ranged from 101–171 g AFDM m⁻² across the four study reaches, with ∼4 × higher values in pools compared to erosional habitats. Total benthic organic matter (BOM) values did not change appreciably with season, but coarse particulate organic matter (CPOM, >1 mm) generally decreased and very fine particulate organic matter (VFPOM, 1.6–250 μm) generally increased during the dry season. Average annual seston concentrations ranged from 0.2–0.6 mg AFDM l⁻¹ (fine seston, <754 μm >250 μm) and 2.0–4.7 mg AFDM l⁻¹ (very fine, <250 μm >1.6 μm), with very fine particles composing 85–92% of total seston. Quality of fine seston particles in the two reaches where tadpoles were present was significantly higher (lower C/N) than the two where tadpoles had been severely reduced (P = 0.0028), suggesting that ongoing amphibian declines in this region are negatively influencing the quality of particles exported from headwaters. Compared to forested streams in other regions, these systems receive relatively high amounts of allochthonous litter inputs but have low in-stream storage. Handling editor: J. Padisak     

Resolving the influence of lignin on soil organic matter decomposition with mechanistic models and continental-scale data

Confidence in model estimates of soil CO₂ flux depends on assumptions regarding fundamental mechanisms that control the decomposition of litter and soil organic carbon (SOC). Multiple hypotheses have been proposed to explain the role of lignin, an abundant and complex biopolymer that may limit decomposition. We tested competing mechanisms using data-model fusion with modified versions of the CN-SIM model and a 571-day laboratory incubation dataset where decomposition of litter, lignin, and SOC was measured across 80 soil samples from the National Ecological Observatory Network. We found that lignin decomposition consistently decreased over time in 65 samples, whereas in the other 15 samples, lignin decomposition subsequently increased. These “lagged-peak” samples can be predicted by low soil pH, high extractable Mn, and fungal community composition as measured by ITS PC2 (the second principal component of an ordination of fungal ITS amplicon sequences). The highest-performing model incorporated soil biogeochemical factors and daily dynamics of substrate availability (labile bulk litter:lignin) that jointly represented two hypotheses (C substrate limitation and co-metabolism) previously thought to influence lignin decomposition. In contrast, models representing either hypothesis alone were biased and underestimated cumulative decomposition. Our findings reconcile competing hypotheses of lignin decomposition and suggest the need to precisely represent the role of lignin and consider soil metal and fungal characteristics to accurately estimate decomposition in Earth-system models.     

植物凋落物影响土壤有机质分解的研究进展

植物凋落物是土壤动物和土壤微生物的主要生命物质和能量来源,其类型、组成以及物理化学等性质直接决定了土壤有机质的品质。对植物凋落物的类型、品质、物理性质、层效应和激发效应以及根际碳淀积与土壤有机质分解的关系进行了总结,可为研究植物凋落物对土壤有机质的影响提供理论参考,指出要在全球变暖背景下进一步加强凋落物分解过程中土壤微生物和酶活性变化的研究。     

A non-native invasive grass increases soil carbon flux in a Hawaiian tropical dry forest

Non‐native plants are invading terrestrial ecosystems across the globe, yet little is known about how invasions impact carbon (C) cycling or how these impacts will be influenced by climate change. We quantified the effect of a non‐native C₄ grass invasion on soil C pools and fluxes in a Hawaiian tropical dry forest over 2 years in which annual precipitation was average (Year 1) and ～60% higher than average (Year 2). Work was conducted in a series of forested plots where the grass understory was completely removed (removal plots) or left intact (grass plots) for 3 years before experiment initiation. We hypothesized that grass invasion would: (i) not change total soil C pools, (ii) increase the flux of C into and out of soils, and (iii) increase the sensitivity of soil C flux to variability in precipitation. In grass plots, grasses accounted for 25–34% of litter layer C and ～70% of fine root C. However, no differences were observed between treatments in the size of any soil C pools. Moreover, grass‐derived C constituted a negligible fraction of the large mineral soil C pool (< 3%) despite being present in the system for ≥50 years. Tree litterfall was ～45% lower in grass plots, but grass‐derived litterfall more than compensated for this reduction in both years. Annual cumulative soil‐surface CO₂ efflux (R_soil) was ～40% higher in grass plots in both years, and increased in both treatments by ～36% in the wetter Year 2. Despite minimal grass‐derived mineral soil C, > 75% of R_soil in grass plots was of C₄ (i.e. grass) origin. These results demonstrate that grass invasion in forest ecosystems can increase the flux of C into and out of soils without changing total C pools, at least over the short term and as long as the native tree canopy remains intact, and that invasion‐mediated changes in belowground C cycling are sensitive to precipitation.     

模拟冻融环境下亚高山森林凋落物分解速率及有机碳动态

森林凋落物分解是森林生态系统物质循环的重要环节,季节性冻融交替是影响凋落物分解的重要环境因素之一,但不同林型的凋落物对冻融响应的差异性很少被量化。为了解冻融环境对森林凋落物分解进程的影响,以川西亚高山森林地区阔叶林、针叶林和针阔混交林3种典型林型的凋落物为实验材料,从凋落物基质质量、冻融环境等影响凋落分解的因素着手,采用模拟冻融循环过程（-5-5℃）,研究了冻融循环中3种林型凋落物分解速率和有机碳含量的动态变化。结果发现,3中典型林型凋落物经过不同冻融处理后,其质量损失、质量损失速率均存在显著差异（P<0.05）。混交林凋落物和针叶林凋落物的分解速率呈慢-快-慢的趋势,而阔叶林凋落物的分解速率逐渐减小。在冻融循环处理下,3种林型的凋落物碳绝对含量呈波动下降的趋势,说明微生物固定外源碳和凋落物释放碳间存在动态平衡。相同林型的凋落物在不同冻融处理下,有机碳释放有显著差异（P<0.05）。其中,冻融环境显著（P<0.05）促进了混交林凋落物和针叶林凋落物有机碳的释放,但是对阔叶林凋落物有机碳的释放没有起到促进作用。这表明全球气候变暖情景下,亚高山森林土壤冻融事件频发将加快凋落物的分解,但变化程度受到凋落物质量控制。     

Distinct roles of savanna and forest tree species in regeneration under fire suppression in a Brazilian savanna

Questions: Has fire suppression relaxed barriers to the exchange of species between savanna and forest? Do all species or a subset of species participate in this exchange? Would current vegetation structure persist if fire suppression were to cease? Location: A gallery forest edge in the Cerrado region of central Brazil that burned only once in the past 35 years. Methods: Density of tree seedlings, saplings and adults, leaf area index (LAI), tree basal area and diameter were surveyed in 12, 10 m × 70 m transects centred on and perpendicular to the forest–savanna boundary. Community composition was assessed using non‐metric multi‐dimensional scaling (NMDS). Results: Basal area and LAI declined substantially from forest to savanna, with an associated shift in species composition. Savanna tree species were nearly absent in the forest, but accounted for the majority of stems in the savanna. In contrast, forest species comprised 14% of adults and more than one‐third of juveniles in the savanna. Despite the high diversity of trees (85 species) in the forest, five species play a particularly large role in this initial phase of forest expansion. Reintroduction of fire, however, would result in widespread topkill of juveniles and the majority of adult forest trees, thereby interrupting the succession towards forest. Conclusions: After 35 years during which the site burned only once, the savanna still remains dominated by savanna species. Nevertheless, the dominance of forest juveniles in border and savanna tree communities suggests that with a continued policy of fire suppression, the forest will continue to expand.     

Responses in organic matter accumulation and processing to an experimental wood addition in three headwater streams

1. Additions of large wood are being used to restore streams that have been subjected to channelization, wood removal or riparian timber harvest. This added wood potentially increases channel stability, habitat complexity and organic matter retention and improves habitat and productivity of higher trophic levels. However, few stream restorations monitor restoration effectiveness after project completion. 2. We added 25 aspen logs (each 2.5 m length × 0.5 m diameter) to 100‐m reaches of each of three forested headwater streams in the Upper Peninsula of Michigan, U.S.A. These wood‐poor streams drain forests that were completely harvested of timber over a century ago and have been selectively logged for the past 50–60 years. An upstream unmanipulateds 100‐m reach in each stream served as the control. 3. We evaluated responses in organic matter processing by measuring red maple leaf decomposition 1 year before and 2 years after wood addition. We also quantified coarse organic matter standing stocks in the main channel and in debris accumulations associated with large wood. In response to wood addition, we predicted both organic matter standing stocks and leaf decomposition rates would increase, thereby enhancing resource availability to higher trophic levels. 4. Leaf decomposition rates did not change following wood addition. Temporal variation in rates among streams was mostly explained by differences in degree days, water velocity, scour/burial and water column inorganic nitrogen concentrations, but not large wood. Variation within streams across years was explained by differences in degree days, water velocity and shredder biomass. 5. Contrary to our prediction, organic matter standing stocks did not increase significantly at the reach scale. However, the experimentally added wood retained c. 4% of total annual coarse benthic organic matter (CBOM) in the first year and an additional c. 15% in the second year, suggesting accumulation over time in the manipulated reaches. The CBOM held by the new logs may be more biologically available because it is less susceptible to burial and transport than material in the streambed. 6. Some shredding macroinvertebrates responded to changes caused by the wood additions. In particular, the common caddisfly shredder, Lepidostoma sp., increased in abundance in leaf bags following wood addition, whereas the biomass of the winter stoneflies, Capniidae, declined in the first year. 7. Considerable funds are spent to restore in‐stream habitat, but few restorations are monitored, particularly over long periods (>5 years). Our results show that longer‐term monitoring is needed to determine the efficacy of these restorations on ecosystem function; organic matter decomposition in our low‐gradient streams did not respond to a substantial increase in large wood after 2 years.     

The influence of soil on vegetation structure and plant diversity in different tropical savannic and forest habitats

Aims Soil plays an important role in the formation and heterogeneity of habitats and thus can cause changes in vegetation structure and plant diversity. The differentiation between Cerrado/savanna and forest is well known, but the relationship between soil and habitats from savannic or forest formations still needs to be better understood, particularly in tropical ecotonal areas. We studied the association between attributes of plant communities, namely structure and diversity, and physicochemical characteristics of soils in the Caatinga domain at the transition to Cerrado in Brazil.Methods Chemical and physical analyses of soils were performed in samples of 38 plots from savannic formations and 30 plots from forest formations. Vegetation was characterized floristically and structurally in all plots, five habitats being assessed in each plant formation. Soil features and vegetation parameters were highly distinct among the different habitats.Important findings In general, forest habitats were more nutrient rich than savannic formation. Furthermore, soil variables showed effects both on vegetation structure and on its species diversity, more pronouncedly in the savannic formations. Habitats were structurally distinct, and diversity differed between savannic and forest communities; however, a higher differentiation occurred when the savannic formation habitats were compared among them. Although plant diversity did not differ among forest formation habitats, soil attributes showed a close relationship with edaphic factors and can contribute for similar vegetation. The soil–vegetation relationship in highly diverse ecotonal landscapes is important from the conservation biology point of view and aid in the execution of proactive plans for the maintenance of biodiversity. Thus, we noticed that diversity and soil behaves distinctly between savannic and forest communities.     

Tree species richness in primary and old secondary tropical forest in Singapore

If secondary succession can accumulate species rapidly, then tropical secondary forests may have an important role to play in the conservation of biodiversity. Data on the floristic composition of forest stands in the Central Catchment Nature Reserve, Singapore, have been analysed to investigate the diversity of approximately 100-year-old tropical secondary forest. Classification using TWINSPAN indicated that three floristic communities could be recognized from 59 0.2 ha plots enumerated for trees >30 cm gbh. These were two types of secondary forest, both dominated by Rhodamnia cinerea (Myrtaceae), and dryland primary forest. The secondary forest was developed on land abandoned after use for agriculture at the end of the 19th century. The 16 primary forest plots contained a total of 340 species, more than the 281 recorded from the 43 plots of the two secondary forest types combined. The mean species number per plot in the more diverse of the two secondary forests was only about 60% of the primary forest. Thus the secondary forest, despite a century or so for colonization by species and the presence of contiguous primary forest, was still significantly less diverse than primary forest areas. It is concluded that secondary forest cannot be assumed to accrete biodiversity rapidly in the tropics, and may not be of direct value in conservation. However, other indirect roles, such as providing resources for native animals, and buffering and protecting primary forest fragments may make the protection of secondary forest worthwhile.