Compartmentalization of the soil animal food web as indicated by dual analysis of stable isotope ratios (N/N and C/C)

The soil animal food web has become a focus of recent ecological research but trophic relationships still remain enigmatic for many taxa. Analysis of stable isotope ratios of N and C provides a powerful tool for disentangling food web structure. In this study, animals, roots, soil and litter material from a temperate deciduous forest were analysed. The combined measurement of δ¹⁵N and δ¹³C provided insights into the compartmentalization of the soil animal food web. Leaf litter feeders were separated from animals relying mainly on recent belowground carbon resources and from animals feeding on older carbon. The trophic pathway of leaf litter-feeding species appears to be a dead end, presumably because leaf litter feeders (mainly diplopods and oribatid mites) are unavailable to predators due to large size and/or strong sclerotization. Endogeic earthworms that rely on older carbon also appear to exist in predator-free space. The data suggest that the largest trophic compartment constitutes of ectomycorrhizal feeders and their predators. Additionally, there is a smaller trophic compartment consisting of predators likely feeding on enchytraeids and potentially nematodes.     

Nutrients in litterfall,forest floor and mineral soils in two adjacent forest ecosystems in Greece

The fluxes of masses and the nutrients Ca,Mg,K,N,P and S were determined in the litterfall of two adjacent forest ecosystems of Hungarian oak(Quercus frainetto L.)and European beech(Fagus sylvatica L.)in a mountainous area of northeastern Greece in 2010–2015.The foliar litterfall for both species reached about 70%of the total litterfall,and was significantly higher from the other two fractions(woody and rest litterfall).The fluxes of masses and nutrients were compared between ecosystems for each fraction separately.Only one significant statistical difference was found,that of K in the woody litterfall.In addition,the stocks of masses and nutrients were calculated in the forest floors and mineral soils of the two ecosystems.Likewise,the stocks of nutrients in the forest floors and mineral soils were compared between ecosystems.In the L horizon of the forest floors,statistical differences,as a result of species effect,were found for the stocks of Ca and N.In the FH horizons,the masses and all the nutrient stocks differed significantly,as the beech plot had much higher quantities of organic matter and nutrients.These higher quantities were probably due to low soil temperatures(microclimate)and high acidity in the beech plot(species effect)that slowed down decomposition.In the mineral soils,the propagation of random error derived from random errors of the individual soil layers was an important factor in the statistical comparisons.Because of the soil acidity in the beech plot,the stocks of exchangeable base cations were significantly higher in the oak plot,whereas the other nutrient stocks did not differ.     

Tree identity surpasses tree diversity in affecting the community structure of oribatid mites (Oribatida) of deciduous temperate forests

The role of tree diversity and identity as determinants of soil animal community structure is little understood. In a mature deciduous forest dominated by beech we identified clusters of one, two and three tree species of beech, ash and lime allowing to investigate the role of tree species diversity and identity on the density and community structure of oribatid mites. To relate oribatid mite community structure to environmental factors we measured leaf litter input, fine root biomass, mass of organic layers, topsoil pH and C and N content. We expected oribatid mite density to increase with increasing tree diversity, but we expected the effects of tree species identity to override effects of tree diversity. Further, we hypothesized the density of oribatid mites to be reduced by the presence of beech but increased by the presence of lime and ash. As expected tree diversity little affected oribatid mite communities, whereas tree species identity strongly altered density and community structure of oribatid mites. However, in contrast to our expectations the density of oribatid mites was highest in presence of beech indicating that many oribatid mite species benefit from the presence of recalcitrant litter forming thick organic layers. Especially Oppioidea benefited from the presence of beech presumably due to an increased availability of food resources such as fungi and nematodes. Lower density of oribatid mites in monospecific clusters of lime and ash suggests that oribatid mites did not benefit from high quality litter of these species. Notably, large and strongly sclerotized oribatid mite species, such as Steganacarus magnus and Chamobates voigtsi, benefited from the presence of ash and lime. Presumably, these large species better resist harsh microclimatic conditions in shallow organic layers.     

Leaf-fall phenology along a topography-mediated environmental gradient in a cool–temperate deciduous broad-leaved forest in Japan

Leaf-fall phenology was studied in a cool–temperate deciduous broad-leaved forest in central Japan in relation to the topographic environmental gradients that occur along a short mountain slope. Leaf-fall phenology was described quantitatively using data from leaf litter collected along the slope. In autumn, leaf fall at the study site tended to occur slightly earlier on the upper slope than on the lower slope. This pattern was found at both the stand and the species levels. Our results suggest that leaf-fall phenology may be affected by difference in microclimatic conditions, because environmental conditions are thought to be more severe on the upper slope than on the lower slope. The less intensive methods used in this study, the litter trap method, and Dixon’s model succeeded in quantifying the phenological patterns of leaf fall within stands and within species along the short mountain slope.     

Assemblages of wood-inhabiting fungi related to silvicultural management intensity in beech forests in southern Germany

Current silvicultural treatments in beech forests are aimed at achieving thick logs without discoloured hardwood. Therefore intensive thinning is applied already in younger stands with the objective of large-sized trunks at an age of 100 years. However, this approach bears the risk that dead wood structures and broken trees are completely removed from the forest. The impact of three different silvicultural management intensity levels on wood-inhabiting fungi over decades was investigated in a large beech forest (>10,000 ha) in southern Germany in 69 sampling plots: A Intensive Thinning and Logging with high-value trees, B Conservation-Oriented Logging with integration of special structures such as dead wood and broken trees and C Strict Forest Reserves with no logging for 30 years. The analysis of community showed marked differences in the fungus species composition of the three treatments, independent of stand age. The relative frequencies of species between treatments were statistically different. Indicator species for naturalness were more abundant at sites with low silvicultural management intensity. Fomes fomentarius, the most common fungus in virgin forests and strict forest reserves, is almost missing in forests with high-management intensity. The species richness seemed to be lower where intensive thinning was applied (P = 0.051). Species characteristic for coarse woody debris were associated to low management intensity, whereas species with a significant preference for stumps became more frequent with increasing management intensity. A total amount of dead wood higher than 60 m³/ha was found to enable significantly higher numbers of species indicators of naturalness (P = 0.013). In conclusion, when applying intensive silvicultural treatment, the role of dead wood needs to be actively considered in order to maintain the natural biocoenosis of beech forests.     

Negligible contribution from roots to soil-borne phospholipid fatty acid fungal biomarkers 18:2ω6,9 and 18:1ω9

The phospholipid fatty acid biomarkers 18:1ω9, 18:2ω6,9 and 18:3ω3,6,9 are commonly used as fungal biomarkers in soils. They have, however, also been found to occur in plant tissues, such as roots. Thus, the use of these PLFAs as fungal biomarkers in sieved soil, which may still contain small remains of roots, has been questioned. We used data from a recent beech tree girdling experiment to calculate the contribution of roots to these biomarkers and were able to demonstrate that not more than 0.61% of 18:1ω9 and 18:2ω6,9 in sieved soil samples originated from roots (but 4% of 18:3ω3,6,9). Additionally, the abundance of the biomarker 18:2ω6,9 in the soil was found to be highly correlated to ectomycorrhizal root colonization, which further corroborates its fungal origin. PLFA biomarkers were substantially reduced in vital roots from girdled trees compared to roots of control trees (by up to 76%), indicating that the major part of PLFAs measured in roots may actually originate from ectomycorrhizal fungi growing inside the roots. We calculated, that even a near to 50% reduction in fine root biomass - as observed in the girdling treatment - accounted for only 0.8% of the measured decrease of 18:2ω6,9. Our results demonstrate that both 18:1ω9 and 18:2ω6,9 are suitable biomarkers for detecting fungal dynamics in soils and that especially 18:2ω6,9 is a reliable biomarker to study mycorrhizal dynamics in beech forests.     

Does forest type and vegetation patchiness influence horizontal distribution of soil Collembola in two neighbouring forest sites?

The influence of forest type and heterogeneity of understory vegetation on the horizontal distribution of soil living Collembola was studied in two neighbouring mountain forest sites—a 180-year old beech forest and a 70-year old natural spruce forest. Four homogenous patches with different understory vegetation were chosen within each forest site and sampled 12 times between 1997 and 1998. A total of 56 collembolan species were identified, 51 in the beech forest and 48 in the spruce site. Twenty-three species were rare—they were recorded with low constancy and density. Although both forest sites differed in soil type, humus form and soil chemical parameters, the species composition of their collembolan communities was quite similar (77% shared species). Nevertheless, soil collembolan communities of both forest sites were clearly delimited on both qualitative (presence–absence) and quantitative (in density of individual species) levels as well as in terms of total collembolan density. Mean collembolan density reached 26 650–44 030 ind/m² in the beech patches compared to 44 470–68 050 ind/m² found in the spruce patches. Considerably higher densities of several species in one forest site may indicate more suitable habitat. In spite of similar species composition and minor differences in qualitative parameters among different vegetation patches within one forest site, there was clear variation in density of individual species, particularly between patches with and without herb vegetation. This could reflect different microclimatic conditions, additional litter supply from herbs or indirect interactions of Collembola with their roots.     

Microbial biomass and respiratory activity in soil aggregates of different sizes from three beechwood sites on a basalt hill

We studied the effects of aggregates of different sizes on the soil microbial biomass. The distribution of aggregate size classes (<2, 2–4, 4–10, >10 mm) in the upper mineral soil horizon (A_h layer) was very different in three sites (upper, intermediate, lower) in a beechwood (Fagus sylvatica) on a basalt hill (Germany). Aggregates of different sizes (<2, 2–4, 4–10 mm) contained different amounts of C and N but the C:N ratios were similar. C and N contents were generally higher in smaller aggregates. The maximum initial respiratory response by microorganisms in intact aggregates and in aggregates passed through a 1-mm sieve declined with the aggregate size, but the difference was more pronounced in intact aggregates. Disruption of aggregates generally increased this response, particularly in 4- to 10-mm aggregates in the lower site. Basal respiration differed strongly among sites, but was similar in each of the aggregate size classes. Aggregate size did not significantly affect the specific respiration (g O₂ g^–1 microbial C h^–1) nor the microbial: organic C ratio, but these parameters differed among sites. Microbial growth was increased strongly by passing the soil through a 1-mm sieve in each of the aggregate materials. The growth of microorganisms in disrupted aggregates was similar, and the effect of aggregate disruption depended on the growth of microorganisms in intact aggregates.     

Coarse particulate organic matter is the primary source of mineral N in the topsoil of three beech forests

Forest soils contain a variable amount of organic N roughly repartitioned among particles of different size, microbial biomass and associated with mineral compounds. All pools are alimented by annual litter fall as main input of organic N to the forest floor. Litter N is further subject to mineralization/stabilization recognized as the crucial process for the turnover of litter N. Although it is well documented that different soil types have different soil N stocks, it is presently unknown how different soil types affect the turnover of recent litter N. Here, we compared the potential mineralization of the total soil organic N with that of recent litter-released N in three beech forests varying in their soil properties. Highly ¹⁵N-labelled beech litter was applied to stands located at Aubure, Ebrach, Collelongo, which differ in humus type, soil type and soil chemistry. After 4-5 years of litter decomposition, the upper 3 cm of the organo-mineral A horizon was sampled and the net N mineralization was measured over 112 days under controlled conditions. The origin of mineralized N (litter N versus soil organic N) was calculated using ¹⁵N labeling. In addition, soils were fractionated according to their particle size (>2000 μm, 200-2000 μm, 50-200 μm, <50 μm) and particulate organic matter (POM) was separated from the mineral fraction in size classes, except the <50 μm fraction. Between 41 and 69% of soil organic N was recovered as POM. Litter-released ¹⁵N was mainly to be found in the coarse POM fractions >200 μm. On a soil mass basis, N mineralization was two-fold higher at Aubure and Collelongo than at Ebrach, but, on a soil N basis, N mineralization was the lowest at Collelongo and the highest at Ebrach. On a soil N (or ¹⁵N) basis, mineralization of litter ¹⁵N was two to four-fold higher than mineralization of the average soil N. Furthermore, the δ¹⁵N of the mineral N produced was closer to that of POM than to that of the mineral-bound fraction (<50 μm). Highest rates of ¹⁵N mineralization happened in the soil with the lowest N content, and we found a negative relationship between accumulations of N in the upper A horizon and the mineralization of ¹⁵N from the litter. Our results show that mineral N is preferentially mineralized from POM in the upper organo-mineral soil irrespective of the soil chemistry and that the turnover rate of litter N is faster in soils with a low N content.     

Response of Fagus sylvatica Seedlings to Root Trenching of Overstorey Picea abies

In a trenching experiment the impact of root competition by overstorey Norway spruce on growth and allocation of above-ground woody biomass of advanced planted beech was investigated in two stands over two growing seasons after trenching. Measurements of soil water potential were taken inside and outside trenched areas by tensiometers at 16 measuring points in one stand from May to September 1999. Height and diameter of the seedlings, as well as branch length and diameter, were measured of beech growing in trenched and untrenched conditions in September before trenching (1998) and afterwards (1999 and 2000). Tensiometers inside the trenches reached the expiring value of -0.085 Mpa 30-40 days later, or never, compared with those in untrenched areas. Whereas relative growth rate (RGR) in height was not affected by trenching, RGR both in diameter and in estimated dry weight of the main stem were substantially enhanced in the first growing season after trenching (1999). A different result was obtained in 2000, when RGR in seedling diameter and estimated dry weight of the main stem was slightly reduced by trenching. The only variable that showed increased RGR of the trenched seedlings in both years was the estimated dry weight of the branches, indicating different above-ground biomass allocation patterns of seedlings with and without root competition.