Evidence for facultative deciduousness in Colophospermum mopane in semi‐arid African savannas

Leaf phenology dictates the time available for carbon assimilation, transpiration and nutrient uptake in plants. Understanding the environmental cues that control phenology is therefore vital for predicting climate‐related changes to plant and ecosystem function. In contrast to temperate systems, and to a lesser degree, tropical forest systems, the cues initiating leaf drop in tropical savannas are poorly studied. We investigated the cues for leaf fall in a tropical monodominant arid savanna species, Colophospermum mopane, using an irrigation experiment. We tracked soil moisture, solar radiation, air temperature, leaf water status, leaf health and leaf carbon balance through the dry season in both irrigated and control plants. Water was the primary cue driving leaf loss of C. mopane rather than temperature or light. Trees watered throughout the dry season retained their canopies. These leaves remained functional and continued photosynthesis throughout the dry season. Leaf carbon acquisition rates did not decline with leaf age but were affected by soil moisture availability and temperature. Leaf loss did not occur when leaf carbon gain was zero, or when a particular leaf carbon threshold was reached. Colophospermum mopane is facultatively deciduous as water availability determines leaf drop in this widespread arid savanna species. Obligate deciduosity is not the only successful strategy in climates with a long dry season.     

Patch dieback of Colophospermum mopane in a dysfunctional semi‐arid African savanna

Abstract Patch dieback occurred in an almost monospecific Colophospermum mopane (Kirk ex Benth.) Kirk ex J. Léonard woodland in the Northern Province, South Africa, following severe droughts in 1988–1989 and 1991–1992. Discrete patches of dieback and adjacent paired areas of ‘healthy’ vegetation lost an average of 87 and 13% of basal area to mortality, respectively. Whole trees mostly died on ‘dead’ plots, while single‐stem mortality prevailed on ‘live’ plots. Tree mortality decreased with increasing stem number per tree. Patch dieback did not occur on sandy soils. On fine‐textured soils, variation in soil type, topography or slope did not affect dieback. Dieback was influenced by vegetation structure, soil surface condition and soil chemistry. Intense intertree competition, shown by self‐thinning occurring prior to dieback, was a precondition for dieback. Intertree competition had heightened during the 30 years prior to dieback because of an increase in woody cover. Dieback patches had changed from functioning as sinks of water and sediment to sources of these as a result of loss of perennial herbaceous cover, decreased water retention on bared surfaces, and accelerated erosion during 50 years of livestock ranching. Vegetation had thus become increasingly drought‐prone, exacerbated in places by soils with a high sodium concentration. Dieback had occurred because the water requirements of C. mopane could no longer be met during drought years on the dysfunctional patches.     

川西高原季节性雪被覆盖对窄叶鲜卑花凋落物分解和养分动态的影响

2010年1-5月在川西高原采用人工雪厚度梯度试验(0、30和100 cm),应用网袋分解法对窄叶鲜卑花叶片凋落物进行分解试验,测定了凋落物的分解速率及其养分动态.结果表明:在无雪被覆盖的样地上分解5个月后的凋落物质量损失率为29.9％,而中雪和深雪样地的凋落物质量损失率分别为33.8％和35.2％.分解过程中,凋落物氮存在一定的富集现象,磷处于波动的富集状态,碳质量分数和碳氮比均呈现前期急剧下降后期逐渐上升的趋势.雪被覆盖显著增加了凋落物的质量损失率和氮含量,而对碳和磷含量无显著影响.在川西高原地区,30 cm以上的持续雪被覆盖能够改变凋落物的分解过程,从而可能对土壤营养物质转化和植物群落构建产生实质性的影响.     

干旱区人工防护林带不同林分凋落叶分解及养分释放

2007年10月下旬至2008年11月,采用原位模拟分解网袋法,对新疆克拉玛依市区北郊人工防护林新疆杨、紫穗槐及二者混合凋落叶进行为期365 d的分解及养分释放动态试验.结果表明：树种不同,凋落叶质量损失率的动态变化不同;凋落叶组成对质量损失率有显著影响,与单优林凋落叶相比,紫穗槐与新疆杨凋落叶混合后更易于分解.经修正Olson负指数衰减模型分析,新疆杨凋落叶分解系数最低(k=0.167),混合凋落叶分解系数最高(k=0.275),估测3种凋落叶半分解和95％分解所需时间为2.41～4.19 a和10.79～17.98 a.不同的分解时期3种凋落叶中N、P和K的残留率不同,分解1年后,K为净释放,N和P为固持或从周围环境中吸收而富集.分解过程中,除紫穗槐凋落叶在分解中期有机碳分解率下降外,其他处理凋落叶有机碳分解率均不断上升,1年后分解率在35.5%～44.2%之间.C/N值基本呈下降趋势,分解前期和中期下降幅度较小,后期下降较快.     

Impacts of altered precipitation regimes on soil communities and biogeochemistry in arid and semi‐arid ecosystems

Altered precipitation patterns resulting from climate change will have particularly significant consequences in water‐limited ecosystems, such as arid to semi‐arid ecosystems, where discontinuous inputs of water control biological processes. Given that these ecosystems cover more than a third of Earth's terrestrial surface, it is important to understand how they respond to such alterations. Altered water availability may impact both aboveground and belowground communities and the interactions between these, with potential impacts on ecosystem functioning; however, most studies to date have focused exclusively on vegetation responses to altered precipitation regimes. To synthesize our understanding of potential climate change impacts on dryland ecosystems, we present here a review of current literature that reports the effects of precipitation events and altered precipitation regimes on belowground biota and biogeochemical cycling. Increased precipitation generally increases microbial biomass and fungal:bacterial ratio. Few studies report responses to reduced precipitation but the effects likely counter those of increased precipitation. Altered precipitation regimes have also been found to alter microbial community composition but broader generalizations are difficult to make. Changes in event size and frequency influences invertebrate activity and density with cascading impacts on the soil food web, which will likely impact carbon and nutrient pools. The long‐term implications for biogeochemical cycling are inconclusive but several studies suggest that increased aridity may cause decoupling of carbon and nutrient cycling. We propose a new conceptual framework that incorporates hierarchical biotic responses to individual precipitation events more explicitly, including moderation of microbial activity and biomass by invertebrate grazing, and use this framework to make some predictions on impacts of altered precipitation regimes in terms of event size and frequency as well as mean annual precipitation. While our understanding of dryland ecosystems is improving, there is still a great need for longer term in situ manipulations of precipitation regime to test our model.     

极端干旱区沙土掩埋对凋落物分解速率及盐分含量动态的影响

极端干旱区由于降水稀少, 植被盖度低, 太阳辐射强烈, 以及土壤稳定性差, 导致其凋落物周转不同于非干旱区。为探究极端干旱区凋落物分解规律, 该研究利用凋落物分解袋法, 以塔克拉玛干沙漠南缘沙漠-绿洲过渡带优势物种花花柴(Karelinia caspia)、骆驼刺(Alhagi sparsifolia)和胡杨(Populus euphratica)凋落叶为研究对象, 设置不同的沙土掩埋处理: 地表、2 cm和15 cm埋深, 以模拟自然条件下凋落物分解环境, 测定分解过程中凋落物质量和水溶性盐的变化特征。结果表明: 极端干旱区凋落物分解速率与凋落物初始碳(C)含量、氮(N)含量、C:N和木质素含量的关系与非干旱区存在较大差异, 在地表处理下, 木质素含量越高, 质量损失越快。不同分解环境下凋落物质量和水溶性盐损失具有显著差异, 与15 cm埋深相比, 地表和2 cm埋深处理显著增加了凋落物的质量损失和水溶性盐总量损失。地表处理增加了凋落物分解前期的水溶性盐溶解量。该研究表明, 极端干旱区凋落物分解的驱动机制具有独特性, 由于降水稀少, 土壤微生物的活性较低, 掩埋深度不是驱动凋落物分解的主要因素, 极端干旱区凋落物的分解主要受其他非生物过程如太阳光辐射的影响。     

Functional diversity of macroinvertebrates on abandoned cattle enclosures in a semi‐arid African savannah

Zimbabwe is experiencing a shift in land‐use, away from livestock farming and towards wildlife conservation. The abandonment of livestock farming may have unforeseen consequences on biodiversity and ecosystem functioning, as cattle kraals create valuable nutrient‐rich patches across the semi‐arid savannah. It is unclear how macroinvertebrates functionally respond to such nutrient‐rich patches in semi‐arid savannahs. We analysed functional diversity of both aboveground and belowground taxa on abandoned cattle kraals and savannah control plots in Save valley Conservancy (SVC). We used distance‐based multivariate techniques to estimate indices of functional diversity. Our results indicated that after two decades of abandonment, kraals had higher functional richness (FRic), functional divergence (FDiv) and functional dispersion (FDis) of macroinvertebrates when both aboveground and belowground species are combined. When aboveground macroinvertebrates were considered alone, no difference was observed for all the considered functional indices. However, only FRic was higher on kraals when belowground macroinvertebrates were separately considered. Our results suggest that two‐decade‐old abandoned kraals may have recovered enough for aboveground species to match the surrounding savannah plot and even surpassed the savannah control for belowground species functional diversity.     

Drought conditions alter litter decomposition and nutrient release of litter types in an agroforestry system of China

Evaluating how decomposition rates and litter nutrient release of different litter types respond to changes in water conditions is crucial for understanding global carbon and nutrient cycling. However, it is unclear how decreasing water affects litter mixture interactions for the maize–poplar system in arid regions. Here, the responses of the litter decomposition process and litter mixture interactions in the agroforestry system to changes in water conditions (control, light drought, and moderate drought) were tested. Moderate drought significantly decreased the decomposition rate for poplar leaf and mixed litters, and decomposition rate was significantly reduced for maize straw litter in light and moderate drought stress. The mass loss rates of maize straw and mixed litters were significantly higher than that of the poplar leaf litter under drought conditions, but there was no significant difference among the three litter types in the control. There was no interaction between mass loss of the mixed litter in the control and light drought conditions, and the litter mixture interaction showed nonadditive synergistic interactions under moderate drought. In terms of nutrient release, there was also no interaction between litter mixture with nitrogen and carbon, but there was antagonistic interaction with potassium release under the light drought condition. Our results demonstrate that drought conditions can lead to decreasing decomposition rate and strong changes in the litter mixture interactions from additive effects to nonadditive synergistic effects in moderate drought. Moreover, light drought changed the mixture interaction from an additive effect to an antagonistic interaction for potassium release.     

Biotic degradation at night,abiotic degradation at day: positive feedbacks on litter decomposition in drylands

The arid and semi‐arid drylands of the world are increasingly recognized for their role in the terrestrial net carbon dioxide (CO₂) uptake, which depends largely on plant litter decomposition and the subsequent release of CO₂ back to the atmosphere. Observed decomposition rates in drylands are higher than predictions by biogeochemical models, which are traditionally based on microbial (biotic) degradation enabled by precipitation as the main mechanism of litter decomposition. Consequently, recent research in drylands has focused on abiotic mechanisms, mainly photochemical and thermal degradation, but they only partly explain litter decomposition under dry conditions, suggesting the operation of an additional mechanism. Here we show that in the absence of precipitation, absorption of dew and water vapor by litter in the field enables microbial degradation at night. By experimentally manipulating solar irradiance and nighttime air humidity, we estimated that most of the litter CO₂ efflux and decay occurring in the dry season was due to nighttime microbial degradation, with considerable additional contributions from photochemical and thermal degradation during the daytime. In a complementary study, at three sites across the Mediterranean Basin, litter CO₂ efflux was largely explained by litter moisture driving microbial degradation and ultraviolet radiation driving photodegradation. We further observed mutual enhancement of microbial activity and photodegradation at a daily scale. Identifying the interplay of decay mechanisms enhances our understanding of carbon turnover in drylands, which should improve the predictions of the long‐term trend of global carbon sequestration.     

Characteristics of hollows and hollow‐bearing trees in semi‐arid river red gum woodland and potential limitations for hollow‐dependent wildlife

Up to 37 species of the birds and microbats inhabiting inland Australia are dependent on tree cavities for breeding or roosting. The river red gum (Eucalyptus camaldulensis), a well‐known hollow‐bearing tree species, occurs in linear semi‐arid woodland along thousands of kilometres of ephemeral river channels and is the only tree species that provides widespread, aggregated hollow resources across a landscape otherwise dominated by shrublands. Here we assess the type and quantity of hollows available along ephemeral rivers of the MacDonnell Ranges bioregion in central Australia and determine which characteristics of river red gums best predict the abundance and characteristics of different tree hollows, as first steps towards assessing the current availability of hollows in the region. Approximately a third of all river red gums sampled were hollow‐bearing, but individual trees with abundant hollows were rare. Further, 36% of hollows had an entrance ≤ 5 cm, and 37% had entrances which were 6–10 cm in diameter, whereas only 13% of hollows had an entrance diameter > 20 cm suitable for larger hollow‐using species. Large and high hollows only occurred on trees that did not display post‐disturbance resprouting. Trees with multiple and diverse hollows were rare and tended to be in advanced stages of senescence and had larger stems (82.3 ± 3.33 cm) and were taller (14.4 ± 0.53 m) compared to non‐hollow‐bearing trees (23.44 ± 1.68 cm, 8.0 ± 0.34 m). Further research is required to establish whether the current abundance of hollows and diversity of hollow types are limiting to cavity‐dependent wildlife, and to identify any threats to availability of hollows.     

Species limits and phylogenetic relationships of red‐finned cryptic species of the seasonal killifish genus Hypsolebias from the Brazilian semi‐arid Caatinga (Teleostei: Cyprinodontiformes: Rivulidae)

Members of the Hypsolebias antenori species group comprise a diverse clade of morphologically similar seasonal killifishes occurring in a vast region of the semi‐arid savannah of northeastern Brazil. The present paper focuses on an assemblage of three allopatric cryptic species (H. antenori from isolated coastal river drainages, Hypsolebias igneus from the São Francisco River basin and Hypsolebias coamazonicus sp. nov. from the Parnaíba River basin) sharing almost identical colour patterns, including the presence of an orangish red anal fin in males, thus herein named as the red‐finned assemblage. A tree‐based approach using mt‐DNA (cytochrome b) supports delimitation of all three species, but indicates that the red‐finned assemblage is paraphyletic – H. igneus and H. coamazonicus are closely related to Hypsolebias nudiorbitatus, whereas H. antenori is the sister group to a clade comprising all 13 species of the H. antenori group included in the analysis. Morphological characters are useful to diagnose species, but are not informative for most clades highly supported by molecular data. H. coamazonicus is distinguished from all other congeners by the possession of a dark grey or black stripe on the dorsal fin in males. The basal position of H. antenori is related to uplift episodes involving the Araripe‐Borborema plateau during the Miocene, which isolated the coastal area inhabited by H. antenori from the remaining areas of the Caatinga. The sister group relationship between H. igneus and H. coamazonicus is attributed to a past connection between the São Francisco and Paranaíba River until the Tertiary.     

Insect outbreaks alter nutrient dynamics in a southern African savanna: patchy defoliation of Colophospermum mopane savanna by Imbrasia belina larvae

Invertebrate herbivore outbreaks have important impacts on system biogeochemical cycling, but these effects have been poorly documented in African savanna ecosystems. In semi‐arid African savannas, outbreaks of the lepidopteran Imbrasia belina (mopane worm) affect discrete patches of the dominant Colophospermum mopane trees; larvae may completely defoliate trees for up to six weeks during each of the early and late growing seasons. We studied the impact of mopane worm outbreaks on the availability of nitrogen (N), phosphorus (P), and potassium (K) within mopane savanna by quantifying major nutrient pools in defoliated and non‐defoliated savanna patches, including leaves, leaf litter, worm frass, and the soil beneath trees. Within an outbreak area, approximately 44 percent of trees were infested, supporting ~29,000 worms/ha, leading to ~640 kg/ha dry weight frass deposition at 1.4 g of frass/day‐individual (fourth or fifth instar), compared with an average 1645 kg/ha dry weight of leaf on trees most of which should be deposited by litterfall at the end of the growing season. Frass had twofold higher P, 10 percent higher K, but equivalent N content than litter. Taking frass and litter deposition together, the N, P, and K contents added due to the outbreak event at our study site were 0.88, 5.8, and 2.8 times those measured in non‐outbreak patches, a pattern which was reflected in the nutrient contents of soil surfaces beneath defoliated trees. Invertebrate herbivory appears to be an important driver for mopane savanna but has been largely neglected.     

季节性雪被对高山森林凋落物分解的影响

季节性雪被可能对高山森林凋落物分解产生重要影响, 但一直没有深入的研究。该文采用凋落物分解袋法, 于2010-2012年雪被覆盖下几个关键时期(冻结初期、深冻期和融化期)以及生长季节, 研究了川西高山森林代表性树种岷江冷杉(Abies faxoniana)、红桦(Betula albosinensis)、四川红杉(Larix mastersiana)和方枝柏(Sabina saltuaria)凋落叶在不同厚度冬季雪被下的分解动态。经过两年的分解, 不同雪被覆盖下岷江冷杉凋落物分解率为33.98%-39.55%, 红桦为46.49%-48.22%, 四川红杉为42.30%-44.93%, 方枝柏为40.34%-43.84%。相对于无雪被覆盖环境, 厚型雪被覆盖均小幅提高了4种凋落物两年的失重率(1.57%-5.57%)。3个针叶树种(岷江冷杉、四川红杉和方枝柏) Olson凋落物分解系数k均以厚型雪被覆盖最大, 薄型雪被覆盖最小, 而阔叶树种红桦分解系数k则表现为无雪被>薄型雪被>较厚型雪被>厚型雪被>中型雪被。尽管在第二年生长季中雪被对红桦凋落物分解的促进作用不明显, 但雪被覆盖明显促进了两年各个关键时期岷江冷杉、四川红杉和方枝柏凋落物的分解。第一年雪被期凋落物分解对当年分解总量的贡献达42.5%-65.5%, 季节性雪被变化明显改变了凋落物冬季分解格局, 对深冻期凋落物分解过程影响尤为显著。综上所述, 当前气候变化情景下冬季雪被的减少可能减缓该区森林凋落物分解过程, 但相对于易分解的阔叶凋落物, 针叶凋落物的响应特征可能更为强烈。     

外源氮添加对温带荒漠地表凋落物分解及养分释放的影响

采用分解网袋法,在古尔班通古特沙漠南缘设置对照N₀(0 g N·m^-2·a^-1)、N₅(5 g N·m^-2·a^-1)、N₁₀(10 g N·m^-2·a^-1)和N₂₀(20 g N·m^-2·a^-1)4个施N处理,研究外源N添加对多枝柽柳、盐角草及两者混合凋落物分解过程及养分释放的影响,分析氮沉降对荒漠生态系统凋落物分解的影响。结果表明: 各物种凋落物的分解速率存在显著差异,经过345 d的分解,多枝柽柳、盐角草及混合物在不同N处理间的分解速率分别为0.64～0.70、0.84～0.99和0.71～0.81 kg·kg^-1·a^-1。凋落物分解过程中,N、P均表现为养分的直接释放,试验结束时,N₀、N₅、N₁₀和N₂₀处理单种凋落物及其混合物N分别释放60.6%～67.4%、56.7%～62.6%、57.4%～62.3%、46.8%～63.0%,P分别释放51.9%～77.9%、59.9%～74.7%、53.0%～79.9%、52.3%～76.4%。N处理对单种凋落物及其混合物的分解影响不显著,但各种凋落物的养分动态对N添加的响应不同,N处理抑制了盐角草N、P释放及混合凋落物P释放,而对多枝柽柳无影响。在温带荒漠,适量的N输入对凋落物分解速率影响不大,但可能会延缓个别物种养分向土壤系统的归还。     

Decomposition patterns of leaf litter of seven common canopy species in a subtropical forest: N and P dynamics

Litter decomposition, governing nutrient and C cycling, is strongly influenced by the chemical litter quality. In order to determine the interspecific variation in leaf decomposition rates and to understand the chemical basis for such variation, decomposition dynamics of seven common canopy species was investigated over 2year using the litterbag technique in a subtropical evergreen broad-leaved forest on Okinawa Island, Japan. The species studied are representatives of the vegetation in the study area and differed significantly in their chemical litter quality. Dry mass loss at the end of study varied in the order: Distylium racemosum< Quercus miyagii< Rapanea neriifolia< Symplocos confusa< Castanopsis sieboldii< Schima wallichii< Daphniphyllum glaucescens. All species showed a pattern characterized by a rapid initial decomposition followed by lower rates except for D. glaucescenswhich decomposition rate appeared to be rather constant. In the late phase, decomposition rates were correlated positively to initial N and ash contents and negatively to lignin content, lignin:N, C:N, and C:P ratios. The effects of N and lignin content or lignin:N ratio were stronger than other quality parameters. There was a wide range in patterns of N and P concentrations, from a net accumulation to a rapid loss in decomposition. The correlation between N and P release suggests that N and P dynamics may have influenced each other during litter decomposition. Analysis of initial quality for species showed that the C:P ratios were extremely high (range 1639–3811) but the N:P ratios were from 28 to 56, indicating a likely P-limitation for this forest. Our results suggest that P is an important control of litter decomposition and N and P dynamics.     

Combined effects of nitrogen addition and litter manipulation on nutrient resorption of Leymus chinensis in a semi‐arid grassland of northern China

Plant growth in semi‐arid ecosystems is usually severely limited by soil nutrient availability. Alleviation of these resource stresses by fertiliser application and aboveground litter input may affect plant internal nutrient cycling in such regions. We conducted a 4‐year field experiment to investigate the effects of nitrogen (N) addition (10 g N·m^?2·year^?1) and plant litter manipulation on nutrient resorption of Leymus chinensis, the dominant native grass in a semi‐arid grassland in northern China. Although N addition had no clear effects on N and phosphorus (P) resorption efficiencies in leaves and culms, N fertilisation generally decreased leaf N resorption proficiency by 54%, culm N resorption proficiency by 65%. Moreover, N fertilisation increased leaf P resorption proficiency by 13%, culm P resorption proficiency by 20%. Under ambient or enriched N conditions, litter addition reduced N and P resorption proficiencies in both leaves and culms. The response of P resorption proficiency to litter manipulation was more sensitive than N resorption proficiency: P resorption proficiency in leaves and culms decreased strongly with increasing litter amount under both ambient and enriched N conditions. In contrast, N resorption proficiency was not significantly affected by litter addition, except for leaf N resorption proficiency under ambient N conditions. Furthermore, although litter addition caused a general decrease of leaf and culm nutrient resorption efficiencies under both ambient and enriched N conditions, litter addition effects on nutrient resorption efficiency were much weaker than the effects of litter addition on nutrient resorption proficiency. Taken together, our results show that leaf and non‐leaf organs of L. chinensis respond consistently to altered soil N availability. Our study confirms the strong effects of N addition on plant nutrient resorption processes and the potential role of aboveground litter, the most important natural fertiliser in terrestrial ecosystems, in influencing plant internal nutrient cycling.     

环境因素对干旱半干旱区凋落物分解的影响研究进展

凋落物分解是干旱半干旱区重要的生化过程,也是区域内物质周转与能量流动的关键生态环节,主要受气候、凋落物基质质量（简称凋落物质量）和土壤生物群落等因素的综合影响.本文综合评述了非生物因素(温度、降水、光辐射、土壤有机质等)和生物因素(凋落物质量、土壤微生物、种群组成和群落结构等)对干旱半干旱地区凋落物分解的影响的相关研究进展.在诸多影响因素中,降水与光辐射是最重要的限制因素.降水能够在短期内使凋落物分解速率迅速增加,而干旱半干旱区光照强度大、时间长,UV B引起的光矿化效应能较好地解释区域内凋落物分解规律.凋落物质量和群落结构主要受生态系统类型的影响,属于长期效应.今后凋落物生态研究的重点主要为全球气候变化下各环境因素的交互作用,不同尺度下凋落物分解过程与格局的变化,以及多因素交互作用凋落物分解模型的构建等方面.