Effects of rhizospheres on the community composition of Collembola in a temperate forest

Influence of living roots on the community composition of Collembola was investigated in a coniferous forest of Chamaecyparis obtusa. We conducted a buried pot experiment that constructed two different systems for carbon (C) availability. We used two types of pots with or without a C. obtusa seedling. The former pots were used to make a system with soil including living roots (i.e. a system based on root-derived and litter-derived C), while the latter was equivalent to soil system without living roots (i.e. a system based on litter-derived C). After 8 months, we harvested the pots and examined the collembolan community and environmental factors. The presence of living roots affected collembolan abundance and species-specific responses. These changes could be explained in terms of leaf biomass of the seedlings, indicating a possible linkage between above-ground productivity and collembolan community. Given the possibility that root-derived C is associated with aboveground plant mass, we speculate that collembolan community responds to root-derived C. The three dominant species, which are widespread in Japanese temperate forests, were more abundant in the presence of living roots. Moreover they were positively correlated with leaf biomass in the system with a seedling, again suggesting the fundamental importance of living roots for the organisation of the collembolan community.     

Vertical distribution of Collembola (Hexapoda) and their food resources in organic horizons of beech forests

 Micro-samples of the surface organic horizons of 13 beech forests in Belgium were fixed immediately after collection in ethanol. Collembola (6255 animals) were sorted directly from micro-samples in the laboratory using a dissecting microscope, while the litter/soil matrix was analysed semi-quantitatively. The vertical distribution of Collembolan species was studied by correspondence analysis. Gut contents of animals were examined under a light microscope and their composition was compared with that of the matrix. A consistent association was found between the vertical distribution of gut contents and that of food resources in the immediate proximity of animals. Species differed in their feeding habits but most of them ingested a wide spectrum of food items. Plasticity in the food regime according to depth could be demonstrated in members of the Onychiuridae family. Received: 11 January 1999     

Direct and indirect effects of tree diversity drive soil microbial diversity in temperate deciduous forest

We investigated the link between aboveground and belowground diversity in temperate deciduous forest ecosystems. To this end, we determined the effects of the tree species composition on the biomass and composition of the soil microbial community using phospholipid fatty acid (PLFA) profiles in the Hainich National Park, a deciduous mixed forest on loess over limestone in Central-Germany. We investigated the effects of the leaf litter composition on the microbial community, hypothesizing that distinctive leaf litter compositions increase signature PLFAs. In addition, we studied the impact of clay content, pH and nutrient status of the soil on the microbial community in different surface soil layers. Consequently, soil was sampled from depths of 0-5 cm, 5-10 cm and 10-20 cm. Plots with highest leaf litter diversity had the largest total amounts of fatty acids, but only PLFA 16:1ω5, which is a common marker for arbuscular mycorrhizal fungi, was significantly increased. In the uppermost soil layer, the pH explained most of the variance in microbial composition. In the deeper surface soil layers, nutrients such as carbon, nitrogen and phosphorus determined the microbial abundances and composition. Our results suggest that the soil microbial community is mainly indirectly influenced by aboveground diversity. Changes in soil pH or the soil nutrient status that are driven by specific plant traits like leave litter quality drive these indirect changes. Specific direct interactions are most reasonable for mycorrhizal fungi.     

Influence of four soil maintenance practices on Collembola communities in a Mediterranean vineyard

We studied the influence of four soil maintenance practices on Collembola communities in the soil of a Mediterranean vineyard: (a) postemergence herbicide with glyphosate; (b) postemergence and pre-emergence herbicides with glyphosate, terbuthylazine, diuron and oryzalin; (c) natural flora and (d) tillage to a depth of 10–15 cm. Total Collembola abundance, species diversity and species richness significantly varied between the four practices. Notably, the practice using postemergence and pre-emergence herbicides had significantly lower values. Identification of Collembola at species level allowed an interspecies comparison and revealed significant differences for the most common species between the four practices, with each practice being characterized by a different set of species. None of the species were found to be significantly more abundant in the plots treated with postemergence and pre-emergence herbicides.     

Lipid composition of Collembola and their food resources in deciduous forest stands—Implications for feeding strategies

The lipid composition of Collembola and their potential food resources was assessed in three deciduous forest stands, in order to gain insight into food web linkages under field conditions. Fatty acids (FAs) previously assigned as trophic markers in laboratory experiments were used to investigate feeding strategies in situ. As potential food sources soil microbiota and plant debris were characterised by their phospholipid fatty acid (PLFA) composition. Both the amount and the pattern of PLFAs differed between sites and soil depth, in particular the bacterial and plant marker FAs in the upper soil layer. Thus, the availability of resources for micro-detritivores varied due to forest and soil layer. The lipid composition of vital and senescent beech leaves was predominantly influenced by metabolic status and represented a quite homogenous FA resource across forest stands. Comparing Folsomia quadrioculata, Lepidocyrtus lignorum, Neanura muscorum and Pogonognathellus longicornis between the different forests revealed FA profiles to be predominantly affected by site, suggesting a diet shift depending on resources at hand. However, species-specific differences in individual FAs occurred, likely related to feeding strategy and physiological activity. Lipids of Collembola comprised low amounts of bacterial marker FAs, and bacterial consumption may occur to some extent, particularly on Gram-positives. The marker FA for predatory feeding, 20:1ω9, was found in several species, although in low amounts. This contradicts known feeding habits and caution is advisable in using 20:1ω9 as trophic marker. Overall, as indicated by high proportions of oleic (18:1ω9) or linoleic (18:2ω6,9) acid, most species were either plant litter or fungal feeders, with some transitions. The ratio 18:1ω9/18:2ω6,9 is proposed as a tool to distinguish between these two major feeding strategies in Collembola.     

Effects of microsite, water, weeding, and direct seeding on the regeneration of native and alien species within a Hawaiian dry forest preserve

Tropical dry forests are among the most endangered ecosystems in the world in general and in Hawaii in particular. To investigate the regeneration ecology of native and alien dry forest species on the island of Hawaii, we used a factorial experiment with microsite (sub-canopy vs. inter-canopy), water (supplemental vs. ambient), and weeding (alien species removed vs. not removed) treatments, and also seeded six native woody species into each plot at the start of the experiment. At the end of the 21-month study, the biomass of the volunteer native and alien species (i.e. unplanted species consisting mainly of relatively fast-growing shrubs) was nearly three and 13 times that of the seeded species, respectively. The biomass of the native volunteers was greater in the inter-canopy plots, greater for the seeded species in the sub-canopy plots, and did not differ significantly within this treatment for the alien species. Few species survived in the ambient water plots, resulting in greater biomass in the watered plots for all species. There were no significant differences in the biomass of the native species within the weeded vs. non-weeded plots; on the contrary, we found consistently positive correlations between the abundance of the seeded species and the volunteer native and alien species. Thus it may be possible to restore Hawaii's degraded dry forests by manipulating these naturally recruiting species to create microsites favorable for the eventual re-establishment of the endangered native canopy tree flora.     

Effects of Collembola on root properties of two competing ruderal plant species

Plant roots compete for nutrients mineralised by the decomposer community in soil. By affecting microbial biomass and activity Collembola influence the nutrient availability to plants. We investigated the effect of Collembola (Protaphorura fimata Gisin) on growth and competition between of two plant species, Cirsium arvense L (creeping thistle) and Epilobium adnatum Griseb. (square-stemmed willow herb), in a laboratory experiment. Two seedlings of each plant species were planted in rhizotrons either in combination or in monoculture (intra- and interspecific competition). Interspecific competition strongly reduced total biomass of C. arvense whereas E. adnatum suffered most from intraspecific competition. Collembola neither affected the competitive relationship of the two plant species nor shoot and root biomass. Although Collembola did not affect total root biomass they influenced root morphology of both plant species. Roots grew longer and thinner and had more root tips in presence of Collembola. Root elongation is generally ascribed to the exploitation of nutrient rich patches in soil. We hypothesise that changes in root morphology in presence of Collembola are due to Collembola-mediated changes in nutrient availability and distribution.     

From forest to fen: Microarthropod abundance and litter decomposition in a southern Appalachian floodplain/fen complex

Our study compared decomposition and litter microarthropod abundance among five plant communities in a mountain floodplain/fen complex located in the southern Appalachian Mountains, USA. We found that the least disturbed plant communities, red maple in particular, have the quickest decomposition, the greatest number of litter microarthropods, the highest soil organic carbon, and the lowest soil pH. Positive correlations were shown between soil organic carbon and total microarthropods; negative correlations were found between soil pH and total microarthropods. No correlations were found between soil moisture and decomposition or total microarthropod numbers. We conclude that soil characteristics related to disturbance, rather than to the presence of a closed canopy, are the main influences on decomposition and litter microarthropods.     

Soil fauna abundance and diversity in a secondary semi-evergreen forest in Guadeloupe (Lesser Antilles): influence of soil type and dominant tree species

The importance of secondary tropical forests regarding the maintenance of soil fauna abundance and diversity is poorly known. The aims of this study were (1) to describe soil fauna abundance and diversity and (2) to assess the determinants of soil fauna abundance and diversity in two stands of a tropical semi-evergreen secondary forest. Soil macrofauna and microarthropod abundance and soil macrofauna diversity were described at two sites developed on different soils and with different site histories: (1) a natural secondary stand (natural forest) under two dominant tree species, Pisonia subcordata and Bursera simaruba, and (2) a planted secondary forest (planted forest) under three tree species, B. simaruba, Swietenia macrophylla, and Tabebuia heterophylla. The effects of both soil and main tree species’ litter quality were assessed to explain soil fauna abundance and diversity. The abundance of soil macrofauna was significantly higher in the soil under the planted forest, and soil fauna communities were contrasted between the two sites. In the planted forest, a soil-dwelling macrofauna community developed (mainly consisting of the anecic earthworm Polypheretima elongata). In the natural forest, soil macrofauna and microarthropod communities were located at the soil surface. The effect of plant litter quality varied according to each dominant tree species and was superimposed to soil effect. The lowest macrofauna abundance was associated with B. simaruba in the natural forest. T. heterophylla supported a much greater macrofauna community than the two other tree species studied at the same soil, and it appears likely that this is due to the palatability of its leaves compared with the other trees (low lignin, tannins, soluble phenols).     

Effects of long-term litter manipulation on soil carbon,nitrogen, and phosphorus in a temperate deciduous forest

Changes in above-ground litterfall can influence below-ground biogeochemical processes in forests. In order to examine how above-ground litter inputs affect soil carbon (C), nitrogen (N) and phosphorus (P) in a temperate deciduous forest, we studied a 14-year-old small-scale litter manipulation experiment that included control, litter exclusion, and doubled litter addition at a mature Fagus sylvatica L. site. Total organic C (TOC), total N (TN) and total P (TP), total organic P (TOP), bioavailable inorganic P (Pi), microbial C, N and P, soil respiration and fine root biomass were analyzed in the A and in two B horizons. Our results showed that litter manipulation had no significant effect on TOC in the mineral soil. Litter addition increased the bioavailable Pi in the A horizon but had no significant effect on N in the mineral soil. Litter exclusion decreased TN and TP in the B horizon to a depth of 10 cm. In the A horizon of the litter exclusion treatment, TP, TOP and bioavailable Pi were increased, which is most likely due to the higher root biomass in this treatment. The high fine root biomass seems to have counteracted the effects of the excluded aboveground litter. In conclusion, our study indicates that aboveground litter is not an important source for C in the mineral soil and that P recycling from root litter might be more important than from above-ground litter.     

Effect of hydrologic regime and forest age on Collembola in riparian forests

The principal objective of the study was to assess factors of primary importance for collembolan community variability measured in riparian forest stands of different age after traditional clear cutting within natural and altered hydrologic regimes. The study was conducted in the Ukrainian part of the Latorica river floodplain where the largest stands of the highly protected medio-European Querco—Ulmetum minoris fluvial forests can be found. Six oak forest stands were investigated on the river floodplain: three in a natural section (inside levee) of the floodplain, which have their original features preserved with periodical inundation, and three in a section separated by a flood control embankment (outside levee) and subjected to serious damage by drainage. In each section the three stands sampled were represented by different ages regenerated with clear-cutting (>3 years, >8 years and >112 years).Differences in hydrologic regimes were central to identification of a predictable and explainable percentage of variation in species composition of Collembola communities in fluvial forests. The hydrologic regime factor was correlated with the basic structural attributes of collembolan communities, such as abundance, species richness and some ecological traits. Season accounted for a higher degree of variation in collembolan communities than did the age of forest stands. We conclude that differences in hydrologic regime are of much higher importance in structuring collembolan communities in riparian forests than are the ages of stands.     

Interactions between mycorrhizal fungi and Collembola: effects on root structure of competing plant species

Mycorrhizal fungi influence plant nutrition and therefore likely modify competition between plants. By affecting mycorrhiza formation and nutrient availability of plants, Collembola may influence competitive interactions of plant roots. We investigated the effect of Collembola (Protaphorura fimata Gisin), a mycorrhizal fungus (Glomus intraradices Schenck and Smith), and their interaction on plant growth and root structure of two plant species, Lolium perenne L. (perennial ryegrass) and Trifolium repens L. (white clover). In a laboratory experiment, two individuals of each plant species were grown either in monoculture or in competition to the respective other plant species. Overall, L. perenne built up more biomass than T. repens. The clover competed poorly with grass, whereas the L. perenne grew less in presence of conspecifics. In particular, presence of conspecifics in the grass and presence of grass in clover reduced shoot and root biomass, root length, number of root tips, and root volume. Collembola reduced shoot biomass in L. perenne, enhanced root length and number of root tips, but reduced root diameter and volume. The effects of Collembola on T. repens were less pronounced, but Collembola enhanced root length and number of root tips. In contrast to our hypothesis, changes in plant biomass and root structure in the presence of Collembola were not associated with a reduction in mycorrhizal formation. Presumably, Collembola affected root structure via changes in the amount of nutrients available and their spatial distribution.     

Experimental analysis of the effect of exotic and native plant species on the structure and function of soil microbial communities

Invasions of exotic plant species are among the most pervasive and important threats to natural ecosystems, however, the effects of plant invasions on soil processes and the soil biota have rarely been investigated. We grew two exotic and a native under-story plant species in the same mineral soil from a non-invaded forest stand in order to test whether observed differences in the field could be experimentally produced in the greenhouse. We characterized changes in the soil microbial community structure (as indexed by PLFAs) and function (as indexed by enzyme activities and SIR), as well as changes in potential nitrogen mineralization rates. We found that the invasion of two very dissimilar exotic species into the under-story of deciduous forests in eastern North America can rapidly cause changes in most of the studied soil properties. At the end of the three-month incubation, soils under the exotic species had significantly different PLFA, enzyme and SIR profiles than both initial soils and soils where native shrubs had been grown. We also observed a significant increase in pH and nitrification rates under one of the exotic plants. Such changes in the soil are potentially long-term (e.g. changes in soil pH) and are therefore likely to promote the re-invasion of these and other exotics. Both management of exotic plant invasions and the restoration of native communities must take into account exotic species effects on the soil.     

刺萼龙葵入侵对不同类型土壤特性的影响

外来入侵植物对土壤生态系统的影响已成为入侵生态学研究的热点。本文选取河北宣化和吉林白城2个刺萼龙葵(Solanum rostratum Dunal.)入侵典型地区,研究刺萼龙葵入侵对不同类型土壤养分及酶活性的影响。每个地区分别选择4个生境作为样地,宣化地区生境包括荒地、路边、农田和林地,白城地区生境包括荒地、路边、农田和草原,其中宣化的林地和白城的草原为每个地区特有的生境。采用滴定法和比色法测定土壤养分和酶活性。结果表明,与未入侵样地(CK)相比,刺萼龙葵入侵显著提高了农田、林地和草原的土壤全氮含量(P0.05),其中宣化(灌淤土)农田和林地全氮含量比CK增加102.49%、79.02%,白城(淡黑钙土)农田和草原比CK增加44.04%、21.81%;刺萼龙葵入侵显著提高了农田和林地的土壤铵态氮含量(P0.05),其中宣化农田、林地中土壤铵态氮含量比CK增加137.59%、50.32%,白城农田中增加140.18%;刺萼龙葵入侵也提高了路边、林地、农田(白城)和荒地(宣化)的土壤硝态氮含量(P0.05),其中,宣化荒地、路边和林地土壤硝态氮含量较CK增加65.52%、32.96%和22.68%,白城路边和农田分别增加132.59%、153.96%。刺萼龙葵入侵显著提高了宣化地区及白城荒地和农田的土壤有机质含量(P0.05);降低了宣化荒地、农田和林地的土壤全磷含量(P0.05),土壤全磷分别比CK降低54.63%、70.81%和29.68%,白城路边和草原的土壤全磷含量分别较CK下降17.40%、14.69%(P0.05);刺萼龙葵入侵对2个地区土壤全钾和速效钾则无显著影响。刺萼龙葵入侵显著提高了荒地、农田、林地、草原和路边(宣化)的土壤脲酶活性(P0.05);宣化路边和白城荒地、草原的中性磷酸酶的活性,宣化荒地和林地土壤过氧化氢酶活性均显著提高(P0.05),对白城过氧化氢酶则无显著影响。刺萼龙葵入侵对某些生境的土壤养分和酶活性无显著影响可能与入侵生境的小气候、入侵时间长短等因素相关,需进一步深入研究。总之,刺萼龙葵入侵改变了2个地区土壤理化性质和酶活性,其可能通过改变土壤养分及土壤酶活性,以创造对自身有利的土壤环境,从而进一步扩散蔓延。     

Controls of temporal and spatial variability of methane uptake in soils of a temperate deciduous forest with different abundance of European beech (Fagus sylvatica L.)

Aerated forest soils are a significant sink for atmospheric methane (CH₄). Soil properties, local climate and tree species can affect the soil CH₄ sink. A two-year field study was conducted in a deciduous mixed forest in the Hainich National Park in Germany to quantify the sink strength of this forest for atmospheric CH₄ and to determine the key factors that control the seasonal, annual and spatial variability of CH₄ uptake by soils in this forest. Net exchange of CH₄ was measured using closed chambers on 18 plots in three stands exhibiting different beech (Fagus sylvatica L.) abundance and which differed in soil acidity, soil texture, and organic layer thickness. The annual CH₄ uptake ranged from 2.0 to 3.4 kg CH₄-C ha⁻¹. The variation of CH₄ uptake over time could be explained to a large extent (R² = 0.71, P < 0.001) by changes in soil moisture in the upper 5 cm of the mineral soil. Differences of the annual CH₄ uptake between sites were primarily caused by the spatial variability of the soil clay content at a depth of 0-5 cm (R² = 0.5, P < 0.01). The CH₄ uptake during the main growing period (May-September) increased considerably with decreasing precipitation rate. Low CH₄ uptake activity during winter was further reduced by periods with soil frost and snow cover. There was no evidence of a significant effect of soil acidity, soil nutrient availability, thickness of the humus layer or abundance of beech on net-CH₄ uptake in soils in this deciduous forest. The results show that detailed information on the spatial distribution of the clay content in the upper mineral soil is necessary for a reliable larger scale estimate of the CH₄ sink strength in this mixed deciduous forest. The results suggest that climate change will result in increasing CH₄ uptake rates in this region because of the trend to drier summers and warmer winters.     

Sources of spatial and temporal variability of inorganic nitrogen in pure and mixed deciduous temperate forests

We investigated the influence of tree canopy composition and structure on the spatial and temporal variability of (i) concentrations of inorganic N (NH₄⁺ and NO₃⁻) and (ii) net N-mineralization and net nitrification, within the temperate forest floor. We compared a pure European beech stand (PS) with a mixed beech-hornbeam one (MS). Three sampling areas were chosen in each stand. Within the PS, the tree locations represented a decreasing gradient of light intensity reaching the forest floor. Within the MS they represented a gradient in the amount of hornbeam leaves present in the litter. In the field NH₄⁺ and NO₃⁻ concentrations were measured in the upper mineral soil (UMS) and the overlying organic layers (OL and OF+OH). Field exposures using buried bags were carried out on UMS over 1 year to measure in situ net N-mineralization and net nitrification. Potential net N-mineralization and net nitrification were investigated in summer with UMS, OL and OF+OH incubated at 28 °C for 28 days in the laboratory. We hypothesize that with the presence of a mull-forming species (hornbeam) within a stand dominated by a moder-forming one (European beech), (i) the spatial and (ii) temporal patterns of soil inorganic N concentrations, net N-mineralization and net nitrification would be different in the two stands. Our main results show that tree species composition has an influence on both spatial and temporal patterns of nitrification. The PS exhibited its highest peaks of UMS NO₃⁻ concentration and net nitrification in spring and early summer while they were highest in the MS in winter. Furthermore, PS exhibited a higher rate of net nitrification than MS. We discuss this unexpected result and suggest that dissolved organic C may be the controlling factor for net nitrification in the MS.     

Assessment of the species composition of forest floor horizons in mixed spruce-beech stands by Near Infrared Reflectance Spectroscopy (NIRS)

How the mixture of tree species modifies short-term decomposition has been well documented using litterbag studies. However, how litter of different tree species interact in the long-term is obscured by our inability to visually recognize the species identity of residual decomposition products in the two most decomposed layers of the forest floor (i.e. the Oe and Oa layers respectively). To overcome this problem, we used Near Infrared Reflectance Spectroscopy (NIRS) to determine indirectly the species composition of forest floor layers. For this purpose, controlled mixtures of increasing complexity comprising beech and spruce foliage materials at various stages of decomposition from sites differing in soil acid-base status were created. In addition to the controlled mixtures, natural mixtures of litterfall from mixed stands were used to develop prediction models. Following a calibration/validation procedure, the best regression models to predict the actual species proportion from spectral properties were selected for each tree species based on the highest coefficient of determination (R²) and the lowest root mean square error of prediction (RMSEP). For the validation, the R² (predictions versus true proportions) were 0.95 and 0.94 for both beech and spruce components in mixtures of materials at all stages of decomposition from the gradient of sites. The R² decreased only marginally by 0.04 when models were tested on independent samples of similar composition. The best models were used to predict the beech-spruce proportion in Oe and Oa layers of unknown composition. They provided in most cases plausible predictions when compared to the composition of the canopy above the sampling points. Thus, tedious and potentially erroneous hand sorting of forest floor layers may be replaced by the use of NIRS models to determine species composition, even at late stages of decomposition.     

Variations of soluble organic nitrogen and microbial nitrogen in deciduous forest soils along an urban-rural gradient

Soluble organic N and microbial N pools in terrestrial ecosystems have been less studied than those of inorganic N. Therefore, cross-system variation in their pool sizes and seasonal dynamics, both absolute and relative to inorganic N pools, needs to be quantified so that their ecological importance in different ecosystems can be evaluated. We measured extractable soil organic N (ESON), microbial biomass N (MBN), and the net production rates of ESON in red oak-dominated remnant forests, along an urban-rural gradient in the New York City metropolitan area. We were interested in (1) determining the seasonal dynamics of ESON and MBN, and (2) examining whether the contrasts in land use (urban, suburban, rural) surrounding these forest remnants were associated with different amounts of ESON and MBN. This field-based study was conducted continuously for 16 months. Yearly average ESON concentrations ranged from 60 to 140 mg kg⁻¹ soil organic matter (SOM), 3-4 times those of inorganic N, and average MBN ranged from 600 to 1100 mg kg⁻¹ SOM. There was a considerable MBN increase in spring in all plots across the gradient. The average increase expressed on an areal basis (to a depth of 7.5 cm) ranged from 1.75 to 4.19 g N m⁻². The N incorporated into the microbial biomass in spring was gradually released later in the growing season (the mean MBN decrease ranged from 1.11 to 3.82 g N m⁻²). The spring MBN increase could be an important retention mechanism for conserving soil inorganic N when plant N uptake may be low. The amplitude in the seasonal dynamics of ESON was far less than that of inorganic N, as was that of net ESON production rates when compared to net N mineralization. These suggest a closer coupling of plant N uptake with inorganic N, much more than with ESON. Both ESON and MBN were significantly higher in rural soils than in urban soils, and both concentrations were positively correlated with SOM content. The variation in ESON, MBN, or SOM associated with this urbanization gradient suggests that the form of N exported, the plant N budget and soil N retention mechanisms may be differentially affected by urban, suburban and rural land uses.