Multi-scale evaluation of soil functional indicators for the assessment of water and soil retention in Mediterranean semiarid landscapes

Ecosystem monitoring and assessment are often based on functional indicators, which provide integrated and yet simple and affordable measures of key ecosystem functions. The landscape function analysis (LFA) assesses ecosystem functioning through three indices that represent basic soil functions: surface stability, infiltration, and nutrient cycling. Given the high scale-dependency of hydrological and erosion processes in semiarid ecosystems, the validation of the stability and infiltration indices requires a multi-scale approach, which has not been applied by previous works. Using records from a four-year monitoring of a semiarid landscape in SE Spain, we evaluated the LFA infiltration and stability indices against quantitative measurements of water and sediment flows at multiple scales. At the finest scale, the indices correctly reflected the higher infiltration and lower sediment production of plant patches as compared with bare-soil interpatches. The infiltration index also captured the spatial variation in the infiltration capacity of bare-soil interpatches. At the hillslope scale, total runoff was inversely related to the average infiltration index for bare-soil interpatches, but it was not related to the global infiltration index, which combines the values from both bare-soil interpatches and plant patches. These results suggest that the hydrological response of semiarid hillslopes depends mainly on the variation in the functioning of bare-soil interpatches. Total sediment yield from the hillslope plots was not related to the stability index. At the catchment scale, both the bare-soil interpatch and the global infiltration indices correctly captured the variability in total runoff produced by three micro-catchments of comparable size. The bare-soil infiltration index predicted bare-soil infiltration rate and hillslope runoff better than common simple indicators of soil functioning such as soil organic carbon, stone cover, crusted bare-soil cover, bulk-density and plant cover, and exhibited a similarly high indicatory potential that a variety of plant spatial-pattern indicators. In contrast to the multi-scale validation of the infiltration index, the indicatory potential of the stability index was only proved for the most contrasting soil conditions in the study site, pointing to a lower sensitivity of this latter index.     

Organic mound-building ants: their impact on soil properties in temperate and boreal forests

Ants are important components of most soil invertebrate communities, and can affect the flow of energy, nutrients and water through many terrestrial ecosystems. The vast majority of ant species build nests in the mineral soil, but a small group of ants in temperate and boreal forests of Eurasia and North America build large parts of their nests above‐ground using organic materials collected from the surrounding soil. Many studies have shown that ants nesting in mineral soil can affect water infiltration rates, soil organic matter (OM) content, and nutrient cycling, but much less is known on how mound‐building ants influence soil physical and chemical properties. In this paper we summarize what is known on the soil impacts of organic mound‐building ants in temperate and boreal forests, and how these ants could be affected by ecosystem disturbance and future climate change. Much of this information comes from studies on Formica rufa group ants in Europe, which showed that CO₂ emissions and concentrations of C, N, and P are usually higher in ant mounds than in the surrounding forest soil. However, ant mounds are a minor component of total soil C and nutrient pools, but they do increase spatial heterogeneity of soil water and available nutrients. Mound‐building ants can also impact tree growth, which could change the quantity and quality of OM added to soil. Forest management, fire, and projected climate change, especially in boreal forests, could affect mound‐building ant population dynamics, and indirectly, soil properties.     

Spatial variability and abiotic determinants of termite mounds throughout a savanna catchment

Termite mounds contribute to the spatial heterogeneity of ecological processes in many savannas, but the underlying patterns and determinants of mound distributions remain poorly understood. Using the Carnegie Airborne Observatory (CAO), we mapped the distribution of termite mounds across a rainfall gradient within a river catchment (～ 27 000 ha) of the Kruger National Park, South Africa. We assessed how different factors were associated with the distribution and height of termite mounds at three spatial scales: the entire catchment, among three broad vegetation types, and on individual hillslope crests. Abiotic factors such as the underlying geology and mean annual precipitation shaped mound densities at broad scales, while local hillslope morphology strongly influenced mound distribution at finer scales, emphasising the importance of spatial scale when assessing mound densities. Fire return period had no apparent association with mound densities or height. Mound density averaged 0.46 mounds ha^?1, and exhibited a clustered pattern throughout the landscape, occurring at relatively high densities (up to 2 mounds ha^?1) on crests, which are nutrient‐poor elements of the landscape. Mounds exhibited significant over‐dispersion (even spacing) at scales below 60 m so that evenly spaced aggregations of termite mounds are embedded within a landscape of varying mound densities. The tallest mounds were found in dry savanna (500 mm yr^?1) and were positively correlated with mound density, suggesting that dry granitic savannas are ideal habitat for mound‐building termites. Mound activity status also varied significantly across the rainfall gradient, with a higher proportion of active (live) mounds in the drier sites. The differential spacing of mounds across landscapes provides essential nutrient hotspots in crest locations, potentially sustaining species that would otherwise not persist. The contribution to biodiversity and ecosystem functioning that mounds provide is not uniform throughout landscapes, but varies considerably with spatial scale and context.     

Ant‐termite interactions: an important but under‐explored ecological linkage

Animal interactions play an important role in understanding ecological processes. The nature and intensity of these interactions can shape the impacts of organisms on their environment. Because ants and termites, with their high biomass and range of ecological functions, have considerable effects on their environment, the interaction between them is important for ecosystem processes. Although the manner in which ants and termites interact is becoming increasingly well studied, there has been no synthesis to date of the available literature. Here we review and synthesise all existing literature on ant–termite interactions. We infer that ant predation on termites is the most important, most widespread, and most studied type of interaction. Predatory ant species can regulate termite populations and subsequently slow down the decomposition of wood, litter and soil organic matter. As a consequence they also affect plant growth and distribution, nutrient cycling and nutrient availability. Although some ant species are specialised termite predators, there is probably a high level of opportunistic predation by generalist ant species, and hence their impact on ecosystem processes that termites are known to provide varies at the species level. The most fruitful future research direction will be to evaluate the impact of ant–termite predation on broader ecosystem processes. To do this it will be necessary to quantify the efficacy both of particular ant species and of ant communities as a whole in regulating termite populations in different biomes. We envisage that this work will require a combination of methods, including DNA barcoding of ant gut contents along with field observations and exclusion experiments. Such a combined approach is necessary for assessing how this interaction influences entire ecosystems.     

The effects of harvester ant (Messor ebeninus Forel) nests on vegetation and soil properties in a desert dwarf shrub community in north-eastern Arabia

In desert ecosystems, harvester ants have been shown to be important granivores and seed dispersers. Because many desert plants exist as seeds in the ground for long periods, harvester ants may greatly influence plant population dynamics. In this study, we examine the effects of harvester ant nests on vegetation and soil properties in a desert ecosystem dominated by the dwarf shrub Rhanterium epapposum in Kuwait, north-eastern Arabia. Soil properties were greatly modified in the circular refuse zone around the nests of the harvester ants, with elevated levels of nutrients (except nitrogen) and organic matter. Plant species richness and productivity were also significantly enhanced in the refuse zone. The vast majority of species inhabiting this zone were desert annuals, but there is no evidence to suggest that the species involved have any distinct association with ant nests. Harvester ants therefore contribute substantially to small-scale spatial heterogeneity in this aridland plant community. It is also suggested that the significance of ant nests in providing favourable sites for plant growth is enhanced in drier years, when many species in off-nest sites may die prematurely due to drought. Furthermore, because ants occasionally abandon their nests and create new colonies, the addition of nutrient-rich patches to the landscape over time could represent an important mechanism for maintaining fertility of desert soils, possibly with long-term implications for plant biodiversity. Harvester ants can therefore be regarded as a key ecosystem engineer in this relatively undisturbed desert ecosystem of Kuwait.     

黄土高原小流域土壤养分的空间异质性

利用地理信息系统的空间分析功能,通过地统计学的半变异函数定量研究了黄土高原典型小流域土壤养分的空间异质性特征。结果表明;土壤有机质,全氮,有效氮,全磷和有效磷的理论模型均为球状模型,由随机因素引起的空间变异占空间总变异的比例小,其值分别为13.333%,10.938%,22.000%,9.091%和27.536%,反映5种养分具有较强的空间自相关格局,但它们的空间自相关范围具有明显的差异。土壤全氮和有效氮变程小,分别是90m和110m,有机质次之,变程是120m,而土壤全磷和有效磷的变程最大为160m,研究成果将有效地指导土壤的取样设计,以及进行土壤养分的空间内插和制图。     

Leaf‐cutting ants,seasonal burning and nutrient distribution in Cerrado vegetation

Abstract Fire and herbivory are known to modify plant community structure. Many studies have suggested that fire ashes may increase soil nutrients in dystrophic soils. Herbivores may also change plant community structure through direct effects of herbivory and affecting nutrient cycling. Leaf‐cutting ants were traditionally viewed as herbivores, although their role may be more complex, because their nests affect both chemical and physical soil properties, thus affecting plants indirectly. We investigated the effects of frequent burning and of leaf‐cutting ants on the nutrient status of an herbaceous and a shrub species occurring in the Brazilian Cerrado, a habitat that is characterized by natural burnings. The proximity of ant nests resulted in an increase of nutrients in the leaves of both vegetation strata, whereas burning sometimes resulted in a decrease of nutrients. Our results do not lead to a possible positive effect of fire on plant nutrient content. On the other hand, ant nests may represent an important source of nutrients for plants on the nutrient‐depleted Cerrado soils and may accelerate vegetation recovery after burning.     

Ecosystem engineering and predation: the multi-trophic impact of two ant species

1. Ants are ubiquitous ecosystem engineers and generalist predators and are able to affect ecological communities via both pathways. They are likely to influence any other terrestrial arthropod group either directly or indirectly caused by their high abundance and territoriality. 2. We studied the impact of two ant species common in Central Europe, Myrmica rubra and Lasius niger, on an arthropod community. Colony presence and density of these two ant species were manipulated in a field experiment from the start of ant activity in spring to late summer. 3. The experiment revealed a positive influence of the presence of one ant colony on densities of decomposers, herbivores and parasitoids. However, in the case of herbivores and parasitoids, this effect was reversed in the presence of two colonies. 4. Generally, effects of the two ant species were similar with the exception of their effect on Braconidae parasitoid densities that responded positively to one colony of M. rubra but not of L. niger. 5. Spider density was not affected by ant colony manipulation, but species richness of spiders responded positively to ant presence. This effect was independent of ant colony density, but where two colonies were present, spider richness was significantly greater in plots with two M. rubra colonies than in plots with one colony of each ant species. 6. To test whether the positive ecosystem engineering effects were purely caused by modified properties of the soil, we added in an additional experiment (i) the soil from ant nests (without ants) or (ii) unmodified soil or (iii) ant nests (including ants) to experimental plots. Ant nest soil on its own did not have a significant impact on densities of decomposers, herbivores or predators, which were significantly, and positively, affected by the addition of an intact nest. 7. The results suggest an important role of both ant species in the grassland food web, strongly affecting the densities of decomposers, herbivores and higher trophic levels. We discuss how the relative impact via bottom-up and top-down effects of ants depends on nest density, with a relatively greater top-down predatory impact at higher densities.     

Local and landscape‐scale effects of an ant nest construction in an open dry forest of Uruguay

1. Ants modify soil properties via nest construction and by doing this modulate soil resources for other organisms. In this sense ants are recognised as ecosystem engineers. 2. In this framework, two less well‐studied issues are focused on: (i) the permanence of the effects of ant nests on plant communities after colonies have died, and (ii) the scaling up from patch to landscape‐scale effects. 3. The aim of the present study was to address these issues in the open dry forests of Uruguay, inhabited by the ant Atta vollenweideri Forel. The active and abandoned nests of this ant represent different and conspicuous patches (30–60 m²) in the landscape. 4. The soil concentration of sodium, a key element in the system, was substantially higher among active nests, and remains high during the early stages of abandoned nests. Woody species abundance, richness, and composition were affected at the patch scale, and simulation models suggested an increase in species richness at the landscape scale. 5. The present study highlights the importance of abandoned nests for plant‐species richness in the ecosystem engineer framework and the need to advance in an integrative approach to study both local and landscape effects of ant's nests.     

蚂蚁扰动对土壤有机碳循环过程的影响研究进展

蚂蚁作为生态系统的消费者和分解者,其对土壤有机碳库的影响一直是学术界研究的热点。目前研究主要从蚂蚁对土壤宏量元素储量、理化性质、微生物群落活动等方面探究蚂蚁对土壤有机碳库的影响。本文综述了蚂蚁扰动对土壤有机碳循环过程特征的影响。蚂蚁筑巢改变了蚁穴土壤的微生境、微气候与土壤理化性质,并通过重构土壤微生物群落结构特征、调控地表植被演替过程与格局等方式,直接或间接的影响蚁巢中土壤有机碳来源、碳库分配过程、有机碳库稳定性、有机质微观分子特征等,在微域、局地乃至景观尺度上影响土壤有机碳的循环过程。未来研究应着重从量化蚂蚁扰动及其导致的环境因子波动对土壤碳通量变化的贡献、建立定量模型联系并统一蚂蚁影响下土壤碳循环过程、厘清蚂蚁影响土壤有机碳库稳定性的机制等方面开展深入研究,揭示蚂蚁作为“生态工程师”在调控土壤碳循环过程中的作用机制。     

Early Response of Soil Properties and Function to Riparian Rainforest Restoration

Reforestation of riparian zones is increasingly practiced in many regions for purposes of biodiversity conservation, bank stabilisation, and improvement in water quality. This is in spite of the actual benefits of reforestation for recovering underlying soil properties and function remaining poorly understood. Here we compare remnant riparian rainforest, pasture and reforestation plantings aged 2–20 years in an Australian subtropical catchment on ferrosols to determine the extent to which reforestation restores key soil properties. Of the nine soil attributes measured (total nitrogen, nitrate and ammonium concentrations, net nitrification and ammonification rates, organic carbon, bulk density, fine root biomass and water infiltration rates), only infiltration rates were significantly lower in pasture than remnant riparian rainforest. Within reforestation plantings, bulk density decreased up to 1.4-fold and infiltration rates increased up to 60-fold with time post-reforestation. Our results suggest that the main outcome of belowground processes of early reforestation is the recovery of the soils'' physical structure, with potential beneficial ecosystem services including reduced runoff, erosion and associated sediment and nutrient loads in waterways. We also demonstrate differential impacts of two commonly planted tree species on a subset of soil properties suggesting that preferential planting of select species could accelerate progress on specific restoration objectives.     

The influence of ants on soil and water losses from an orange orchard in eastern Spain

Herbicide applications have greatly reduced plant cover, and increased soil erosion on a new orange orchard planted on valley slopes in eastern Spain. This has increased the importance of soil fauna, such as ants, in regulating soil erosion processes. Ants increase water infiltration rates by forming soil macropores during nest construction, but new soil brought to the surface by ant activity could increase the sediments available for erosion. Simulated rainfall experiments were conducted on 20 paired plots (20 with ant activity and 20 controls) to study the impact of ants on surface water flow and sediment movement in an intensively managed orange orchard near Valencia, Spain. Simulated rainfall was applied to each plot at a rate of 55 mm/h on a 0.25 m² area for 1 h. We found a reduction of soil bulk density, an increase in soil organic matter, and an increase in macropore flow in ant‐affected soils, as compared to soil without ant activity. These ant‐induced soil changes increased water infiltration rates and runoff discharge. However, the fresh, unconsolidated soil brought to the surface during nest construction resulted in greater soil loss on two plots than their ant‐free controls. Ants can be an important factor in soil erosion processes when surface vegetation is removed by intensive herbicide use.     

Landscape evolution and nutrient rejuvenation reflected in Amazon forest canopy chemistry

Terra firme forests make up more than three quarters of the western Amazon basin and are often considered functionally homogeneous in regional scale mapping and modelling efforts. However, the landforms underlying these systems are subject to dynamic processes of landscape evolution occurring within an otherwise geomorphically stable terrace formation. These processes may introduce systematic variability in local nutrient status of terra firme ecosystems. We utilised high‐resolution airborne topographic and imaging spectroscopy data, with directed field soil surveys, to reveal that active stream incision and patterns of soil rock derived nutrient availability drive foliar canopy chemistry distributions across seven catchments within a single terrace formation. These results strongly suggest that fine‐scale geomorphic processes directly affect biogeochemical cycles throughout the lowland western Amazon. Furthermore, links between landscape evolution and foliar chemical distributions indicate that geomorphic processes drive the fine‐scale spatial organisation of this tropical ecosystem, with implications for the functional assembly and biogeography of Amazonian forests.     

南京东郊城市湿地水质多尺度空间分异

对南京市东郊的17个城市湿地进行水质监测和植物群落调查,从生态系统、集水区和景观功能区3个尺度揭示城市湿地水质的分异特征.结果表明:湿地类型和植物群落特征是生态系统尺度上影响湿地水质的主要因素;原生湿地的水质普遍优于改造湿地,而改造湿地的水质则普遍优于次生湿地;湿地植物的盖度越大,水质越好.土地利用类型是集水区尺度上影响湿地水质空间分异的主要因素,建设用地对流域湿地营养盐影响程度较大,而林地对溶解氧影响程度较大.人类活动是功能区尺度上影响湿地水质空间分异的主要因素,而其对不同水质指标的影响程度不同.     

Influence of an invasive ant on grazing and detrital communities and nutrient fluxes in a tropical forest

Aim  Pathways linking grazing and detrital subsystems of terrestrial ecosystems are important for ecosystem processes and function, but remain poorly understood. The invasion of a generalist predator creates a unique opportunity to study the effects of predation across these subsystems. We examine here, the effects of a non-native generalist predator, the little red fire ant ( Wasmannia auropunctata, Roger) on both grazing and detrital invertebrate communities and ecosystem processes in a rain forest understorey.
Location  Gamba Protected Area Complex, south-western Gabon, Africa.
Methods  We measured abundances and diversities of understorey grazing and detrital invertebrate communities, soil nutrients, herbivory, litter fragmentation rates and leaf chemistry of a dominant understorey shrub inside and outside of 19 separate invasion fronts. We then explored possible trophic cascades and pathways of interaction using path analysis.
Results  Results suggest that invasive ants may alter herbivory regimes, grazing and detrital communities, and may indirectly alter litter decomposition and nutrient cycling in the soil by suppressing important microbivore and detritivore populations with consequences for leaf chemistry.
Main conclusions  These results demonstrate that generalist predators may be major drivers of both grazing and detrital subsystems by inducing strong shifts in adjacent communities that ultimately affect ecosystem processes.     

Indicators of landscape organization and functionality in semi-arid former agricultural lands under a passive restoration management over two periods of abandonment

Abandoned lands previously used for agricultural purposes may constitute an opportunity for understanding the variables involved in the restoration of native ecosystems over time. In this study, we assessed the functional status of an abandoned farmland currently used for conservation, using a methodology based on indicators of landscape organization and soil surface. We analyzed changes in plant cover, patch and interpatches structure and several soil surface properties during two periods of land abandonment (less and more than 40 years). Using this methodology, we characterize the potential of the ecosystem to capture and transfer resources, and the state of functional properties such as infiltration capacity, soil stability and nutrient cycling.We detected a significant development of the shrub layer and an increase of the number, type and area occupied by vegetated patches in old fields. The contribution of vegetated patches to the recovery of stability, infiltration and nutrient functions was significant at the local scale. Nonetheless, when analyzing the landscape as a whole only the nutrient cycling index was significantly higher in old fields. The lack of improvement of the stability and infiltration in old fields can lead to further degradation and indicates that even though vegetation cover may have improved over time in old fields, the current cover might not be enough to prevent further degradation by erosion. Our results reinforce the importance of using functionality indexes in future studies focused on adequate restoration measures to protect the function of desert ecosystems, especially when aimed for biodiversity conservation. Hence, the use of monitoring systems based on organization and function indicators is a useful tool to represent the current state and potential recovery of previously disturbed ecosystems, provided that the time scale is taken into account.     

Invasional 'meltdown' on an oceanic island

Islands can serve as model systems for understanding how biological invasions affect community structure and ecosystem function. Here we show invasion by the alien crazy ant Anoplolepis gracilipes causes a rapid, catastrophic shift in the rain forest ecosystem of a tropical oceanic island, affecting at least three trophic levels. In invaded areas, crazy ants extirpate the red land crab, the dominant endemic consumer on the forest floor. In doing so, crazy ants indirectly release seedling recruitment, enhance species richness of seedlings, and slow litter breakdown. In the forest canopy, new associations between this invasive ant and honeydew‐secreting scale insects accelerate and diversify impacts. Sustained high densities of foraging ants on canopy trees result in high population densities of host‐generalist scale insects and growth of sooty moulds, leading to canopy dieback and even deaths of canopy trees. The indirect fallout from the displacement of a native ‘keystone’ species by an ant invader, itself abetted by introduced/cryptogenic mutualists, produces synergism in impacts to precipitate invasional ‘meltdown’ in this system.     

Litter Biomass and Nutrient Determinants of Ant Density, Nest Size, and Growth in a Costa Rican Tropical Wet Forest

The nutritional demands of animals vary by taxon. Across landscapes, communities of animals experience variability in the stoichiometry of carbon and nutrients within their resource base. Thus, we expect stoichiometry to contribute to the spatial variance in the demographic parameters of animal communities. Here, we measure how the composition of a litter-nesting tropical rainforest ant community is influenced by spatial variation in environmental stoichiometry relative to litter biomass, a known predictor of ant density. We found the density of ants and their nests were strongly related to litter biomass and carbon: phosphorus stoichiometry. The spatial variation in soil nutrients, which determines leaf litter stoichiometry, was an excellent predictor of nest size in the two most common genera of ants. We found a negative relationship between species' growth rate and local soil stocks of phosphorus. Overall, the density of litter-dwelling ants varied greatly across this tropical forest landscape and environmental stoichiometry can account for limits on ant density independent of the biomass of the leaf litter resource base.     

植物与土壤微生物在调控生态系统养分循环中的作用

陆地生态系统的地上、地下是相互联系的。植物与土壤微生物作为陆地生态系统中的重要组成部分, 它们之间的相互作用是生态系统地上、地下结合的重要纽带。该文首先介绍了植物在养分循环中对营养元素的吸收、积累和归还等作用, 阐述了土壤微生物对养分有效性及土壤质量具有重要的作用。其次, 重点综述了植物与土壤微生物之间相互依存、相互竞争的关系。植物通过其凋落物与分泌物为土壤微生物提供营养, 土壤微生物作为分解者提供植物可吸收的营养元素, 比如共生体菌根真菌即可使植物根与土壤真菌达到互惠。然而, 植物的养分吸收与微生物的养分固持同时存在, 因而两者之间存在对养分的竞争。通过植物多样性对土壤微生物多样性的影响分析, 以及土壤微生物直接或间接作用于植物多样性和生产力的分析, 探讨了植物物种多样性与土壤微生物多样性之间的内在联系。针对当前植物与土壤微生物对养分循环的调控机制的争论, 提出植物凋落物是调节植物与土壤微生物养分循环的良好媒介, 植物与土壤微生物的共同作用对维持整个生态系统的稳定性具有重要意义。也指出了目前在陆地生态系统地上、地下研究中存在的不足和亟待解决的问题。     

The Role of Wood Ants (Formica rufa group) in Carbon and Nutrient Dynamics of a Boreal Norway Spruce Forest Ecosystem

Wood ants (Formica rufa group) are regarded as keystone species in boreal and mountain forests of Europe and Asia by their effect on ecosystem carbon (C) and nutrient pools and fluxes. To quantify the impact of their activity on boreal forest ecosystems, C, nitrogen (N), phosphorus (P), potassium (K) and calcium (Ca) pools and fluxes in wood ant nests (WAN), and soil were assessed along a 5-, 30-, 60-, and 100-year-old Norway spruce (Picea abies L. Karsten) dominated successional gradient in eastern Finland. Amounts of C and nutrients in WAN increased with stand age, but contained less than 1% of total C and nutrient pools in these stands. The CO₂-efflux from nests was also insignificant, as compared to CO₂-efflux from the forest floor. Annually, the amount of C brought by wood ants into their nests as honeydew, prey and nest-building materials ranged from 2.7 to 49.3 kg ha^?1 C, but this is only 0.1–0.7% of the combined net primary production of trees and understorey in boreal forests. The difference between wood ant nest C inputs and outputs was very small in the younger-aged stands, and increased in the older stands. Carbon accumulation rates in nests over a 100 year period are estimated to be less than 10 kg ha^?1 a^?1. In contrast to C, annual inputs of N, P, and K are larger compared to wood ant nest nutrient pool size, ranging from 3 to 6% of the annual tree stand and understorey uptake. This indicates a more rapid turnover and transport of N, P, and K out of WAN, and suggests that wood ants increase the cycling rate of these nutrients in boreal forests.