Disentangling the Litter Quality and Soil Microbial Contribution to Leaf and Fine Root Litter Decomposition Responses to Reduced Rainfall

Climate change-induced rainfall reductions in Mediterranean forests negatively affect the decomposition of plant litter through decreased soil moisture. However, the indirect effects of reduced precipitation on litter decomposition through changes in litter quality and soil microbial communities are poorly studied. This is especially the case for fine root litter, which contributes importantly to forests plant biomass. Here we analyzed the effects of long-term (11 years) rainfall exclusion (29% reduction) on leaf and fine root litter quality, soil microbial biomass, and microbial community-level physiological profiles in a Mediterranean holm oak forest. Additionally, we reciprocally transplanted soils and litter among the control and reduced rainfall treatments in the laboratory, and analyzed litter decomposition and its responses to a simulated extreme drought event. The decreased soil microbial biomass and altered physiological profiles with reduced rainfall promoted lower fine root—but not leaf—litter decomposition. Both leaf and root litter, from the reduced rainfall treatment, decomposed faster than those from the control treatment. The impact of the extreme drought event on fine root litter decomposition was higher in soils from the control treatment compared to soils subjected to long-term rainfall exclusion. Our results suggest contrasting mechanisms driving drought indirect effects on above-(for example, changes in litter quality) and belowground (for example, shifts in soil microbial community) litter decomposition, even within a single tree species. Quantifying the contribution of these mechanisms relative to the direct soil moisture-effect is critical for an accurate integration of litter decomposition into ecosystem carbon dynamics in Mediterranean forests under climate change.     

Cross‐ecosystem effects of a large terrestrial herbivore on stream ecosystem functioning

Large terrestrial consumers have direct and indirect effects on forest ecosystem function, but few studies have investigated the impacts of terrestrial consumers on freshwater ecosystems. In the Cape Breton Highlands of Nova Scotia, browsing by hyper‐abundant moose following a spruce budworm outbreak has transformed boreal forest into grasslands. We conducted a field study to investigate the potential for cross‐ecosystem effects of hyper‐abundant moose following budworm outbreak on small boreal stream ecosystem structure and function. With our field study, we tested the prediction that watersheds with higher levels of moose‐mediated grasslands in their sub‐basin would have higher stream temperatures, total nitrogen, electrical conductivity, periphyton biomass and macroinvertebrate abundances. While our data supported several of our predictions pertaining to moose impacts on the abiotic variables (i.e. temperature range, total nitrogen, electrical conductivity) we found evidence of variable moose impacts on the benthic community. Specifically, we observed lower relative abundance of predatory invertebrates in streams with high moose impacts compared to streams with low moose impacts in their watersheds but no evidence of moose impacts on the relative abundance of shredders, filterers, gatherers, and grazers. This empirical study fills a key gap in our understanding of spatial ecosystem ecology by providing insight into the effects of large terrestrial consumers across ecosystem boundaries with potential implications for landscape‐scale management of hyper‐abundant ungulates. Given the broad availability and improvement in remote sensing technology, the novel integration of remote sensing and field studies employed here may provide a roadmap for future studies of meta‐ecosystem dynamics.     

Plant growth response to direct and indirect temperature effects varies by vegetation type and elevation in a subarctic tundra

There has been growing recent use of elevational gradients as tools for assessing effects of temperature changes on vegetation properties, because these gradients enable temperature effects to be considered over larger spatial and temporal scales than is possible through conventional experiments. While many studies have explored the direct effects of temperature, the indirect effects of temperature through its long‐term influence on soil abiotic or biotic properties remain essentially unexplored. We performed two climate chamber experiments using soils from a subarctic elevational gradient in Abisko, Sweden to investigate the direct effects of temperature, and indirect effects of temperature via soil legacies, on growth of two grass species. The soils were collected from each of two vegetation types (heath, dominated by dwarf shrubs, and meadow, dominated by graminoids and herbs) at each of three elevations. We found that plants responded to both the direct effect of temperature and its indirect effect via soil legacies, and that direct and indirect effects were largely decoupled. Vegetation type was a major determinant of plant responses to both the direct and indirect effects of temperature; responses to soils from increasing elevation were stronger and showed a more linear decline for meadow than for heath soils. The influence of soil biota on plant growth was independent of elevation, with a positive influence across all elevations regardless of soil origin for meadow soils but not for heath soils. Taken together, this means that responses of plant growth to soil legacy effects of temperature across the elevational gradient were driven primarily by soil abiotic, and not biotic, factors. These findings emphasize that vegetation type is a strong determinant of how temperature variation across elevational gradients impacts on plant growth, and highlight the need for considering both direct and indirect effects of temperature on plant responses to future climate change.     

Spatial patterns of total and available N and P at alpine treeline

Background and aims

Vegetation can have direct and indirect effects on soil nutrients. To test the effects of trees on soils, we examined the patterns of soil nutrients and nutrient ratios at two spatial scales: at sites spanning the alpine tundra/subalpine forest ecotone (ecotone scale), and beneath and beyond individual tree canopies within the transitional krummholz zone (tree scale).

Methods

Soils were collected and analyzed for total carbon (C), nitrogen (N), and phosphorus (P) as well as available N and P on Niwot Ridge in the Colorado Rocky Mountains.

Results

Total C, N, and P were higher in the krummholz zone than the forest or tundra. Available P was also greatest in the krummholz zone while available N increased from the forest to the tundra. Throughout the krummholz zone, total soil nutrients and available P were higher downwind compared to upwind of trees.

Conclusions

The krummholz zone in general, and downwind of krummholz trees in particular, are zones of nutrient accumulation. This pattern indicates that the indirect effects of trees on soils are more important than the direct effects. The higher N:P ratios in the tundra suggest nutrient dynamics differ from the lower elevation sites. We propose that evaluating soil N and P simultaneously in soils may provide a robust assay of ecosystem nutrient limitation.     

Phosphorus Limitation of Microbial Processes in Moist Tropical Forests: Evidence from Short-term Laboratory Incubations and Field Studies

Although there is a widespread belief that phosphorus (P) limits basic ecosystem processes in moist tropical forests, direct tests of this supposition are rare. At the same time, it is generally believed that P does not limit soil microorganism respiration or growth in terrestrial ecosystems. We used natural gradients in P fertility created by soils of varying age underlying tropical rain forests in southwestern Costa Rica, combined with direct manipulations of carbon (C) and P supply, to test the effects of P availability on the decomposition of multiple forms of C, including dissolved organic carbon (DOC) and soil organic carbon (SOC). Results from a combination of laboratory and field experiments suggest that C decomposition in old, highly weathered oxisol soils is strongly constrained by P availability. In addition, P additions to these soils (no C added) also revealed that microbial utilization of at least labile fractions of SOC was also P limited. To our knowledge, this is the first direct evidence of P limitation of microbial processes in tropical rain forest soil. We suggest that P limitation of microbial decomposition may have profound implications for C cycling in moist tropical forests, including their potential response to increasing atmospheric carbon dioxide. Furthermore, this site is still relatively rich in P when compared to many other tropical forests on old soils; thus, we believe that P limitation of soil microorganisms throughout the humid tropics is a possibility.     

增温和隔离降雨对亚热带森林土壤N2O通量的影响

设置对照(CT)、增温5 ℃(W)、隔离50%降雨(P)和增温5 ℃+隔离50%降雨(WP)4种处理,以相关功能基因作为标志物,研究增温和隔离降雨影响亚热带森林生态系统土壤N₂O通量变化的途径.结果表明: 隔离降雨显著降低了土壤铵态氮浓度;增温显著降低了土壤N₂O通量和土壤反硝化势.增温处理(W)和降雨处理(P)的土壤微生物生物量氮(MBN)均显著低于对照(CT),AOA amoA基因丰度与MBN和铵态氮含量之间呈显著负相关,但与土壤硝化势和土壤N₂O通量没有显著相关性.路径分析显示,反硝化势直接显著影响土壤N₂O通量,而微生物生物量磷(MBP)和增温则通过直接影响反硝化势来间接影响土壤N₂O通量.温度可能是影响亚热带森林土壤N₂O通量的主要驱动因素,全球变暖可能会减少亚热带森林土壤的N₂O排放.     

增温和隔离降雨对亚热带森林土壤N2O通量的影响

设置对照(CT)、增温5 ℃(W)、隔离50%降雨(P)和增温5 ℃+隔离50%降雨(WP)4种处理,以相关功能基因作为标志物,研究增温和隔离降雨影响亚热带森林生态系统土壤N₂O通量变化的途径.结果表明: 隔离降雨显著降低了土壤铵态氮浓度;增温显著降低了土壤N₂O通量和土壤反硝化势.增温处理(W)和降雨处理(P)的土壤微生物生物量氮(MBN)均显著低于对照(CT),AOA amoA基因丰度与MBN和铵态氮含量之间呈显著负相关,但与土壤硝化势和土壤N₂O通量没有显著相关性.路径分析显示,反硝化势直接显著影响土壤N₂O通量,而微生物生物量磷(MBP)和增温则通过直接影响反硝化势来间接影响土壤N₂O通量.温度可能是影响亚热带森林土壤N₂O通量的主要驱动因素,全球变暖可能会减少亚热带森林土壤的N₂O排放.     

Amazonian Dark Earth Shapes the Understory Plant Community in a Bolivian Forest

Amazonian Dark Earths (ADE) are the result of human modification of the Amazonian landscape since pre‐Columbian times. ADE are characterized by increased soil fertility compared to natural soils. In the Amazonian forest, soil fertility influences understory herb and fern species composition. However, little research has been done to evaluate the effect of ADE on the composition of the understory community. We evaluated the effects of ADE and soil in 36 plots (150 m × 4 m) established in a Bolivian moist forest (La Chonta). For each plot, we determined soil nutrients, and the composition, richness, and abundance of terrestrial fern, angiosperm herb, and understory palm species. We found that the presence of ADE created a gradient in soil nutrients and pH that affected the understory species composition especially of ferns and palms. Additionally, the higher nutrient concentration and more neutral soil pH on ADE soils caused a decrease of ferns species richness. We therefore conclude that the current composition of the understory community in this particular Bolivian forest is a reflection of past human modifications of the soil.     

Spatial patterns of herbivory by gall-forming insects: a test of the soil fertility hypothesis in a Mexican tropical dry forest

It has been proposed that fertile soils reduce the incidence of gall-forming insect (GFI) species in plant communities. This is known as the soil fertility hypothesis. The main objective of this study was to analyze the spatial distribution of GFI species under different habitats in a tropical dry forest at the Chamela-Cuixmala Biosphere Reserve, Mexico. Eight habitats which differ in soil type, topography, nutrient availability and vegetation were chosen. We found that 38 GFI species specialize on their host plant species. GFI species richness was negatively correlated with phosphorous and nitrogen availability. Using phosphorous as an indicator of soil fertility, we found low frequency and density of GFI on fertile soils. Our study indicates that soil fertility is one of the factors that negatively affects the patterns of spatial distribution of species richness, incidence and abundance of GFI at the community level in two different ways: i) indirectly affecting GFI species richness in plants adapted to infertile soils and ii) directly affecting GFI responses to plant traits of hosts found in a fertility gradient.     

Soil biota reduce allelopathic effects of the invasive Eupatorium adenophorum

Allelopathy has been hypothesized to play a role in exotic plant invasions, and study of this process can improve our understanding of how direct and indirect plant interactions influence plant community organization and ecosystem functioning. However, allelopathic effects can be highly conditional. For example allelopathic effects demonstrated in vivo can be difficult to demonstrate in field soils. Here we tested phytotoxicity of Eupatorium adenophorum (croftonweed), one of the most destructive exotic species in China, to a native plant species Brassica rapa both in sand and in native soil. Our results suggested that natural soils from different invaded habitats alleviated or eliminated the efficacy of potential allelochemicals relative to sand cultures. When that soil is sterilized, the allelopathic effects returned; suggesting that soil biota were responsible for the reduced phytotoxicity in natural soils. Neither of the two allelopathic compounds (9-Oxo-10,11-dehydroageraphorone and 9b-Hydroxyageraphorone) of E. adenophorum could be found in natural soils infested by the invader, and when those compounds were added to the soils as leachates, they showed substantial degradation after 24 hours in natural soils but not in sand. Our findings emphasize that soil biota can reduce the allelopathic effects of invaders on other plants, and therefore can reduce community invasibility. These results also suggest that soil biota may have stronger or weaker effects on allelopathic interactions depending on how allelochemicals are delivered.     

Species richness and soil properties in Pinus ponderosa forests: A structural equation modeling analysis

Question: How are the effects of mineral soil properties on understory plant species richness propagated through a network of processes involving the forest overstory, soil organic matter, soil nitrogen, and understory plant abundance? Location: North‐central Arizona, USA. Methods: We sampled 75 0.05‐ha plots across a broad soil gradient in a Pinus ponderosa (ponderosa pine) forest ecosystem. We evaluated multivariate models of plant species richness using structural equation modeling. Results: Richness was highest at intermediate levels of understory plant cover, suggesting that both colonization success and competitive exclusion can limit richness in this system. We did not detect a reciprocal positive effect of richness on plant cover. Richness was strongly related to soil nitrogen in the model, with evidence for both a direct negative effect and an indirect non‐linear relationship mediated through understory plant cover. Soil organic matter appeared to have a positive influence on understory richness that was independent of soil nitrogen. Richness was lowest where the forest overstory was densest, which can be explained through indirect effects on soil organic matter, soil nitrogen and understory cover. Finally, model results suggest a variety of direct and indirect processes whereby mineral soil properties can influence richness. Conclusions: Understory plant species richness and plant cover in P. ponderosa forests appear to be significantly influenced by soil organic matter and nitrogen, which are, in turn, related to overstory density and composition and mineral soil properties. Thus, soil properties can impose direct and indirect constraints on local species diversity in ponderosa pine forests.     

Linking dominant Hawaiian tree species to understory development in recovering pastures via impacts on soils and litter

Large areas of tropical forest have been cleared and planted with exotic grass species for use as cattle pasture. These often remain persistent grasslands after grazer removal, which is problematic for restoring native forest communities. It is often hoped that remnant and/or planted trees can jump‐start forest succession; however, there is little mechanistic information on how different canopy species affect community trajectories. To investigate this, I surveyed understory communities, exotic grass biomass, standing litter pools, and soil properties under two dominant canopy trees—Metrosideros polymorpha (‘ōhi‘a) and Acacia koa (koa)—in recovering Hawaiian forests. I then used structural equation models (SEMs) to elucidate direct and indirect effects of trees on native understory. Native understory communities developed under ‘ōhi‘a, which had larger standing litter pools, lower soil nitrogen, and lower exotic grass biomass than koa. This pattern was variable, potentially due to historical site differences and/or distance to intact forest. Koa, in contrast, showed little understory development. Instead, data suggest that increased soil nitrogen under koa leads to high grass biomass that stalls native recruitment. SEMs suggested that indirect effects of trees via litter and soils were as or more important than direct effects for determining native cover. It is suggested that diverse plantings which incorporate species that have high carbon to nitrogen ratios may help ameliorate the negative indirect effects of koa on natural understory regeneration.     

Surface soil chemical and physical patterns in a brigalow–Dawson gum forest,central Queensland

Brigalow (Acacia harpophylla)–Dawson gum (Eucalyptus cambageana) open forests are predominantly supported by solodic soils in central Queensland. This report describes relations between some physical, chemical and morphological properties of surface soils (0–0.10 m) within a virgin brigalow–Dawson gum forest. Soil property gradients were found to radiate horizontally from tree-dominated to non-vegetated areas, indicative of a vegetation-induction process. These reflect the importance of litterfall organic matter in determining soil fertility and soil physical conditions at this site. The decomposition of litterfall is probably responsible for significant increases in soil organic carbon (C), total nitrogen (N) and total sulphur (S) concentrations with increasing vegetation canopy cover. The composition of soil organic matter appears constant across the study area, as similar soil C:N:S ratios were recorded across all vegetation canopy classes. Soil salinity, total phosphorus, mineral nitrogen, cation exchange capacity, exchangeable calcium and exchangeable potassium levels also increased with increasing vegetation canopy cover. Surface soil physical properties were also related to vegetation canopy cover. Bulk density and the < 20 μm dispersion ratio decreased while soil micro-relief and A hor?izon depth increased with increasing vegetation canopy cover. Organic C, through inverse relations with bulk density and the < 20 μm dispersion ratio, appears to enhance soil porosity and aggregate stability, indicating the role of organic matter in the stabilization of larger virgin soil aggregates. The close association between virgin forest and surface soil fertility is seen to have implications for changes in land-use, with a decline in nutrient availability, soil aggregate stability and productivity following forest clearing.     

Recovery of soil productivity with forest succession on abandoned agricultural land

Much of the primary forest in the eastern United States that was converted to farmland between 1600 and 1900 has reverted back to second growth forest as a result of agriculture abandonment. This reversion back to forest gives soil productivity a chance to recover, though the rates of recovery are not well understood. Understanding the legacy effects of past disturbances like agriculture can provide important insights to support ecological restoration efforts on disturbed soils. Our goal with this study was to further understand the effects of forest development on soil productivity after agriculture abandonment. We used a chronosequence approach to examine soil properties over a 60‐year temporal scale of forest development on abandoned agricultural lands in Saratoga and Rensselaer Counties in New York, U.S.A. We measured soil properties within this chronosequence to test the hypothesis that there would be measurable recoveries of soil physical properties and fertility over time. We observed rapid recovery of physical properties (lower bulk density and higher macroporosity) of surface soils within 5–10 years after agricultural abandonment. However, we found a legacy effect of agricultural compaction still evident in subsoils, with soil strength measurements indicating that past agricultural practices still limited root growth 55–60 years after abandonment. Soil percent organic matter and mineralizable nitrogen (N) both increased with forest development, but biomass accumulation may be slowed by limited root growth in the subsoil due to high strength. We recommend assessing subsoil physical properties when developing ecological restoration plans for agricultural lands.     

森林土壤氮素转换及其对氮沉降的响应

近几十年人类活动向大气中排放的含氮化合物激增 ,并引起大气氮沉降也成比例增加。目前 ,氮沉降的增加使一些森林生态系统结构和功能发生改变 ,甚至衰退。近 2 0 a欧洲和北美有关氮沉降及其对森林生态系统的影响方面的研究较多 ,而我国少有涉及。森林土壤氮素转换是森林生态系统氮素循环的一个重要的组成部分 ,而矿化、硝化和反硝化作用是其核心过程 ,氮沉降作为驱动因子势必改变森林土壤氮素转换速度、方向和通量。根据国外近 2 0 a有关研究 ,首先介绍了森林土壤氮素转换过程和强度 ,论述森林土壤氮素在生态系统氮素循环中的作用 ,然后在此基础上 ,介绍了氮沉降对森林土壤氮素循环的研究途径 ,探讨了氮沉降对森林土壤氮素矿化、硝化和反硝化作用的影响及其机理     

No short-term change in soil properties following four-fold litter addition in a Costa Rican rain forest

We experimentally manipulated forest floor litter to investigate the influence of litter quality and quantity on soil properties over the short-term (weeks to months) in a wet tropical forest in NE Costa Rica. The study included old growth forest on high fertility soils, old growth forest on low fertility soils, and secondary forest on intermediate fertility soils. Forest floor litter was removed from a 16 m² area and added to an adjacent 4 m² area in March 2003, resulting in a one to four-fold increase in the annual litter input to the forest floor. We created three addition, three removal and three control plots per forest type. We measured treatment effects on variation in soil moisture, temperature, pH, and Bray-1 P (plant available) over a 5-month period that captured the dry-wet season transition. Litter manipulation had no effect on any of the soil properties measured during the 5-month study period. Significant variability through time and a similar temporal pattern across the three forest stands suggest that climatic variability is driving short-term patterns in these soil properties rather than seasonal inputs of litter. In general, soils were warmer, drier and more basic with higher available P during dry season months. Even in wet tropical forests, small variability in climate can play an important role in soil dynamics over periods of weeks to months. Although litter manipulation did not influence soil properties over the 5-month study period, a longer lag may exist between the timing of litter inputs and the influence of that litter on soil properties, especially plant available P.     

Microbial resource allocation for phosphatase synthesis reflects the availability of inorganic phosphorus across various soils

According to the resource allocation model for extracellular enzyme synthesis, microorganisms should preferentially allocate their resources to phosphorus (P)-acquiring enzyme synthesis when P availability is low in soils. However, the validity of this model across different soil types and soils differing in their microbial community composition has not been well demonstrated. Here we investigated whether the resource allocation model for phosphatase synthesis is applicable across different soil types (Andosols, Acrisols, Cambisols, and Fluvisols) and land uses (arable and forest), and we examined which soil test P and/or P fraction microorganisms responded to when investing their resources in phosphatase synthesis in the soils. The ratio of alkaline phosphatase (ALP) to β-d-glucosidase (BG) activities in the arable soils and the ratio of acid phosphatase (ACP) to BG activities in the forest soils were significantly negatively related with the available inorganic P concentration. We also observed significant effects of available inorganic P, pH, soil types, and land uses on the (ACP + ALP)/BG ratio when the data for the arable and forest soils were combined and used in a stepwise multiple regression analysis. These results suggest that microbial resource allocation for phosphatase synthesis is primarily controlled by available inorganic P concentration and soil pH, but the effects of soil types and land uses are also significant.     

Soil Effects on Forest Structure and Diversity in a Moist and a Dry Tropical Forest

Soil characteristics are important drivers of variation in wet tropical forest structure and diversity, but few studies have evaluated these relationships in drier forest types. Using tree and soil data from 48 and 32 1 ha plots, respectively, in a Bolivian moist and dry forest, we asked how soil conditions affect forest structure and diversity within each of the two forest types. After correcting for spatial effects, soil‐vegetation relationships differed between the dry and the moist forest, being strongest in the dry forest. Furthermore, we hypothesized that soil nutrients would play a more important role in the moist forest than in the dry forest because vegetation in the moist forest is less constrained by water availability and thus can show its full potential response to soil fertility. However, contrary to our expectations, we found that soil fertility explained a larger number of forest variables in the dry forest (50 percent) than in the moist forest (17 percent). Shannon diversity declined with soil fertility at both sites, probably because the most dominant, shade‐tolerant species strongly increased in abundance as soil fertility increased.     

A review of the role of fire in nutrient cycling in Australian native forests,and of methodology for studying the fire-nutrient interaction*

Previous studies relating to the effects of fire on nutrient cycling in Australian native forests are reviewed. It is apparent that: (a) the action of fire on soil fertility is likely to be important in affecting the functioning of Australian plant communities, (b) work to date only gives a guide as to the nature and magnitude of the effects of fire on some nutrient pools and nutrient cycling processes, but has not established the significance of such effects to the nutrient budget of entire forest systems, or established the links between fire, soil fertility and the functioning of forest communities. The difficulties associated with studying and predicting the long-term effects of fire on the fertility of forest soils are discussed. A new methodology applicable to study of the interaction between fire and plant nutrition is advocated, and is based on examination of the effects of fire on nutrient stores and flux rates in the soil-litter compartment of the forest. This approach emphasizes the use of changes in rates of nutrient cycling processes as an index of the-effects of disturbances such as fire on a forest ecosystem. The application of the methodology in a study of the effects of repeated low-intensity prescribed burning on soil fertility is discussed briefly.     

Tropical forest recovery: legacies of human impact and natural disturbances

Land-use history interacts with natural forces to influence the severity of disturbance events and the rate and nature of recovery processes in tropical forests. Although we are far from an integrated view of forest recovery processes, some generalizations can be made. Recovery of forest structure and composition is relatively rapid following disturbances that primarily impact forest canopies, such as hurricanes. Recovery is considerably slower following disturbances that heavily impact soils as well as aboveground vegetation, such as bulldozing, heavy or long-term grazing, and severe fires, often with long-lasting effects on species composition. The landscape matrix plays a critical role in local recovery processes. Proximity of disturbed areas to remnant forest patches promotes more rapid recovery, which depends heavily on seed dispersal. Recovery of aboveground biomass is constrained by soil fertility and texture across regions as well as across soil types within a region. Restoration of soil fertility may be a prerequisite for forest recovery on sites with severely degraded soils. Despite evidence of rapid forest recovery following large-scale deforestation, many degraded areas of today's tropics will require human assistance to recover forest structure, species composition, and species interactions typical of mature tropical forests.