Effects of biological invasions on forest carbon sequestration

There has been a rapidly developing literature on the effects of some of the major drivers of global change on carbon (C) sequestration, particularly carbon dioxide (CO₂) enrichment, land use change, nitrogen (N) deposition and climate change. However, remarkably little attention has been given to one major global change driver, namely biological invasions. This is despite growing evidence that invasive species can dramatically alter a range of aboveground and belowground ecosystem processes, including those that affect C sequestration. In this review, we assess the evidence for the impacts of biological invaders on forest C stocks and C sequestration by biological invaders. We first present case studies that highlight a range of invader impacts on C sequestration in forest ecosystems, and draw on examples that involve invasive primary producers, decomposers, herbivores, plant pathogens, mutualists and predators. We then develop a conceptual framework for assessing the effects of invasive species on C sequestration impacts more generally, by identifying the features of biological invaders and invaded ecosystems that are thought to most strongly regulate C in forests. Finally we assess the implications of managing invasive species on C sequestration. An important principle that emerges from this review is that the direct effects of invaders on forest C are often smaller and shorter‐term than their indirect effects caused by altered nutrient availability, primary productivity or species composition, all of which regulate long‐term C pools and fluxes. This review provides a conceptual basis for improving our general understanding of biological invaders on ecosystem C, but also points to a paucity of primary data that are needed to determine the quantitative effects of invaders on ecosystem processes that drive C sequestration.     

Soil inorganic carbon storage pattern in China

Soils with pedogenic carbonate cover about 30% (3.44 × 10⁶ km²) of China, mainly across its arid and semiarid regions in the Northwest. Based on the second national soil survey (1979–1992), total soil inorganic carbon (SIC) storage in China was estimated to be 53.3±6.3 PgC (1 Pg=10¹⁵ g) to the depth investigated to 2 m. Soil inorganic carbon storages were 4.6, 10.6, 11.1, and 20.8 Pg for the depth ranges of 0–0.1, 0.1–0.3, 0.3–0.5, and 0.5–1 m, respectively. Stocks for 0.1, 0.3, 0.5, and 1 m of depth accounted for 8.7%, 28.7%, 49.6%, and 88.9% of total SIC, respectively. In contrast with soil organic carbon (SOC) storage, which is highest under 500–800 mm yr⁻¹ of mean precipitation, SIC storage peaks where mean precipitation is <400 mm yr⁻¹. The amount and vertical distribution of SIC was related to climate and land cover type. Content of SIC in each incremental horizon was positively related with mean annual temperature and negatively related with mean annual precipitation, with the magnitude of SIC content across land cover types showing the following order: desert, grassland >shrubland, cropland >marsh, forest, meadow. Densities of SIC increased generally with depth in all ecosystem types with the exception of deserts and marshes where it peaked in intermediate layers (0.1–0.3 m for first and 0.3–0.5 m for latter). Being an abundant component of soil carbon stocks in China, SIC dynamics and the process involved in its accumulation or loss from soils require a better understanding.     

Role of root-mediated interactions in phytotoxic interference of Ageratum conyzoides with rice (Oryza sativa)

Ageratum conyzoides L. (billy goat weed; Asteraceae) is an annual invasive weed native of tropical America and has now naturalized worldwide, particularly in Southeast Asia. The present study investigated the nature and potential of root-mediated allelopathic interference of A. conyzoides against rice (Oryza sativa). Root and shoot length and biomass accumulation of rice were significantly reduced (by 18–30%) when grown in the rhizosphere soil of the weed indicating the release of putative allelochemicals from the weed into the soil. The growth of rice was also progressively reduced in the soil amended with increasing amounts of root residues (5, 10 and 20 g kg⁻¹ soil) of A. conyzoides. The addition of activated charcoal, an inert material with high affinity for organic biomolecules, partly ameliorated the negative effects of root residues amended in the soil. Further, there was no negative effect on the availability of soil nutrients in the root-amended soils. These were rather nutrient rich with greater electrical conductivity, and higher amount of organic matter, thus indicating no role in observed growth reduction. The reduction in allelopathic effects of root residue upon charcoal addition further indicated that putative phytotoxins released from the weed roots are water-soluble phenolic compounds. A significant amount of water-soluble phenolics were present in rhizosphere (∼6-times higher) and root-amended soils (∼5–10-fold higher) and their content was reduced (to ∼3.6–7.0-fold higher) when charcoal was added. The observed growth reduction in Ageratum rhizospheric or root-amended soils was concomitant with the amount of phenolic compounds. Upon HPLC analyses, these were identified as p-coumaric acid, gallic acid, ferulic acid, p-hydroxybenzoic acid and anisic acid. Under laboratory conditions, these phenolic acids reduced the root length and seedling weight of rice individually as well as in equimolar mixture, though no synergistic effect was noticed. The study concludes that root exudates and residues of A. conyzoides suppress the growth of rice by releasing phenolic allelochemicals into the soil rhizosphere and not through alteration of soil nutrients, and allelopathy plays a significant role in root-mediated negative interference of A. conyzoides.     

Tropical nematode diversity: vertical stratification of nematode communities in a Costa Rican humid lowland rainforest

Comparisons of nematode communities among ecosystems have indicated that, unlike many organisms, nematode communities have less diversity in the tropics than in temperate ecosystems. There are, however, few studies of tropical nematode diversity on which to base conclusions of global patterns of diversity. This study reports an attempt to estimate nematode diversity in the lowland tropical rainforest of La Selva Biological Research Station in Costa Rica. We suggest one reason that previous estimates of tropical nematode diversity were low is because habitats above the mineral soil are seldom sampled. As much as 62% of the overall genetic diversity, measured by an 18S ribosomal barcode, existed in litter and understorey habitats and not in soil. A maximum-likelihood tree of barcodes from 360 individual nematodes indicated most major terrestrial nematode lineages were represented in the samples. Estimated 'species' richness ranged from 464 to 502 within the four 40 × 40 m plots. Directed sampling of insects and their associated nematodes produced a second set of barcodes that were not recovered by habitat sampling, yet may constitute a major class of tropical nematode diversity. While the generation of novel nematode barcodes proved relatively easy, their identity remains obscure due to deficiencies in existing taxonomic databases. Specimens of Criconematina, a monophyletic group of soil-dwelling plant-parasitic nematodes were examined in detail to assess the steps necessary for associating barcodes with nominal species. Our results highlight the difficulties associated with studying poorly understood organisms in an understudied ecosystem using a destructive (i.e. barcode) sampling method.     

Long-term CO2 production from deeply weathered soils of a tropical rain forest: evidence for a potential positive feedback to climate warming

Currently, it is unknown what role tropical forest soils will play in the future global carbon cycle under higher temperatures. Many tropical forests grow on deeply weathered soils and although it is generally accepted that soil carbon decomposition increases with higher temperatures, it is not known whether subsurface carbon pools are particularly responsive to increasing soil temperatures. Carbon dioxide (CO₂) diffusing out of soils is an important flux in the global carbon. Although soil CO₂ efflux has been the subject of many studies in recent years, it remains difficult to deduct controls of this flux because of the different sources that produce CO₂ and because potential environmental controls like soil temperature and soil moisture often covary. Here, we report results of a 5‐year study in which we measured soil CO₂ production on two deeply weathered soil types at different depths in an old‐growth tropical wet forest in Costa Rica. Three sites were developed on old river terraces (old alluvium) and the other three were developed on old lava flows (residual). Annual soil CO₂ efflux varied between 2.8–3.6 μmol CO₂‐C m^?2 s^?1 (old alluvium) and 3.4–3.9 μmol CO₂‐C m^?2 s^?1 (residual). More than 75% of the CO₂ was produced in the upper 0.5 m (including litter layer) and less than 7% originated from the soil below 1 m depth. This low contribution was explained by the lack of water stress in this tropical wet forest which has resulted in very low root biomass below 2 m depth. In the top 0.5 m CO₂ production was positively correlated with both temperature and soil moisture; between 0.6 and 2 m depth CO₂ production correlated negatively with soil moisture in one soil and positively with photosynthetically active radiation in the other soil type. Below 2 m soil CO₂ production strongly increased with increasing temperature. In combination with reduced tree growth that has been shown for this ecosystem, this would be a strong positive feedback to ecosystem warming.     

武夷山自然保护区不同海拔甜槠天然林土壤有机碳变化特征及影响因素

为揭示中亚热带常绿阔叶林建群种--甜槠天然林不同海拔土壤有机碳含量垂直分布差异及影响机制，以武夷山自然保护区甜槠天然林单一植被类型为研究对象，在其集中分布的5个海拔梯度（540、700、850、1022、1200 m）范围内设置固定样地，测定每个海拔梯度不同深度土层土壤因子（土壤全氮、全磷、土壤pH值、容重、土壤有机质、粉粒、砂粒、粘粒）、气候因子（土壤温度）、植被因子（细根生物量）及土壤有机碳含量等指标，分析了土壤有机碳沿海拔及垂直土层分布特征，并在主成分分析基础上构建了基于主控因子的线性回归模型。结果表明：（1）同一海拔高度，土壤有机碳含量在土壤垂直剖面分布具有明显的"表聚性"现象；同一土层深度，随着海拔升高，土壤有机碳含量逐渐增加，但增幅随土层深度增加而减小，高海拔地区有助于土壤有机碳的固存；（2）不同土层土壤有机碳含量与海拔、土壤全氮、土壤含水量、土壤粉粒呈极显著正相关（P<0.01），与土壤温度、土壤容重、土壤粘粒、砂粒呈极显著负相关（P<0.01）；土壤细根生物量、土壤有机质与土壤有机碳含量在土壤表层（0-10、10-20 cm）呈极显著（P<0.01）或显著正相关（P<0.05）；土壤pH值、土壤砂粒与土壤有机碳含量在20-30 cm土层呈显著负相关（P<0.05），但与其他土层关系不显著（P>0.05）；海拔因素是影响土壤有机碳含量分布的主要因素，其次为土壤因素，植被因素主要影响土壤表层有机碳含量分布。（3）海拔因素能通过影响与土壤有机碳形成和转化的因子及改变土壤有机碳的累积和分解速率，对土壤有机碳的分布产生影响。（4）多元线性回归模型拟合R²高于一元线性回归模型拟合R²，能解释土壤有机碳含量变异的82.1%-98.1%。由此可见，不同环境因子组合可以更好的解释不同土层土壤有机碳含量随海拔梯度的变异。     

Aerated subsurface irrigation water gives growth and yield benefits to zucchini, vegetable soybean and cotton in heavy clay soils

Inadequate oxygen concentration in the root zone is a constraint to plant performance particularly in heavy, compacted and/or saline soils. Sub-surface drip irrigation (SDI) offers a means of increasing oxygen to plant roots in such soils, provided irrigation water can be hyper-aerated or oxygenated. Hydrogen peroxide (HP) at the rate of 5 litre ha⁻¹ at the end of each irrigation cycle was injected through SDI tape to a field-grown zucchini (courgette) crop (Cucurbita pepo) on a saturated heavy clay soil in Queensland, Australia. Fruit yield, number and shoot weight increased by 25%, 29% and 24% respectively due to HP treatment compared to the control. Two pot experiments with vegetable soybean (Glycine max) and cotton (Gossypium hirsutum) compared the effectiveness of HP and air injection using a Mazzei air injector (a venturi), throughout the irrigation cycle in raising crop yield in a heavy clay soil kept at saturation or just under field capacity. Fresh pod yield of vegetable soybean increased by 82–96% in aeration treatments compared with the control. The yield increase was associated with more pods per plant and greater mean pod weight. Significantly higher above ground biomass and light interception were evident with aeration, irrespective of soil water treatment. Similarly cotton lint yield increased by 14–28% in aeration treatments compared with the control. The higher lint yield was associated with more squares and bolls per plant which accompanied greater above ground biomass and an increase in root mass, root length and soil respiration. Air injection and HP effected greater water use, but also brought about an enhancement of water use efficiency (WUE) for pod and lint yield, and increased leaf photosynthetic rate in both species but had no effect on transpiration rate and stomatal conductance per unit leaf area. Aeration-induced enhanced root function was arguably responsible for greater fruit set and yield in all three crops, while in vegetable soybean greater canopy cover, radiation interception and total vegetative biomass were responsible for additional yield benefit. Increased aeration of the root zone in heavy clay soils employing either air injection or HP proved beneficial to SDI irrigated crops, irrespective of the soil water conditions, and can add value to grower investments in SDI.     

Effects of freeze–thaw on C and N release from soils below different vegetation in a montane system: a laboratory experiment

Unstable snow cover and more frequent freeze–thaw events have been predicted for montane areas in southern Norway, where stable winters are common today. These systems are important contributors to the flux of carbon (C) and nitrogen (N) to air and water. Here we quantify and compare the effects of freeze–thaw on C and N release from soils collected below Calluna, Molinia or Sphagnum. Intact organic soil cores were subjected to four different freeze–thaw regimes for four consecutive 2‐week periods: (1) slow cycling (SC) with one long freezing event during each 2‐week period, (2) fast cycling (FC) with four short freezing events during each 2‐week period, (3) permanent frost (PF) and (4) permanent thaw (PT). The freezing temperature was −5 °C and the thawing temperature was 5 °C. Before start of treatment, at the end of each 2‐week period, and during postincubation periods, carbon dioxide (CO₂) emission as well as leachable dissolved organic C (DOC), dissolved organic N (DON), ammonium (NH₄), nitrate (NO₃) and absorbance at 254 nm were measured. In soils from all three vegetations, PF increased the release of CO₂, DOC, DON and NH₄ compared with PT. SC caused some scattered effects whereas FC only resulted in some increase in NO₃ release below Molinia. Generally, the emission of CO₂ and leaching of DOC, DON and NH₄ increased in the following order: Sphagnum < Calluna < Molinia. The release of NO₃ was greatest below Calluna. Our data suggest that vegetation cover and composition seem at least as important as increased soil frost for future winter fluxes of CO₂, DOC, DON and dissolved inorganic N (DIN) from the soil to air and water. The freezing period needs to be sufficiently long to give significant effects.     

Collembola respond to aphid herbivory but not to honeydew addition

Abstract. 1. Euedaphic collembola alter their soil distribution in response to above‐ground aphid herbivory of Poa annua L. Graminae, a host grass. 2. Two mechanisms potentially underpin this effect. Carbon‐rich aphid honeydew falling onto the soil surface may affect mycophagous collembola; alternatively aphid‐induced changes in root biomass may be necessary to produce changes in collembola abundance. 3. When compared to a plant‐only control, aphid herbivory increased the number of collembola in the top 5 cm of soil, reduced both foliar and root biomass, and increased shoot/root ratio. Honeydew addition had no effect on collembola numbers or any recorded host‐plant parameter. 4. Honeydew deposition is not responsible for the increased numbers of collembola found in the upper soil after aphid herbivory; aphid‐induced reductions in root biomass may be the most important factor explaining knock‐on effects of aphid herbivory on soil fauna.