Changes in microbial nutrient status during secondary succession and its modification by earthworms

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Abstract
Microbial biomass, nutrient (N and P) status, and carbon and nutrient limitation of the microflora were investigated in soils from five different sites (field, 5-, 12-, and about 50-year-old fallow, beechwood), which represent different stages of a secondary succession from a wheat field to the climax ecosystem of a beechwood on limestone. In addition, the effect of faeces production by the substrate feeding earthworm species Octolasion lacteum (Örley) on the nutrient status of the soil microflora of these sites was studied. Humus had accumulated in the soil of the third fallow site, with an enhanced biomass of microflora. However, in the beechwood soil, which had the highest humus content, microbial biomass was lower than in the soil of the third fallow site and similar to that of the field and the two younger fallow sites. In general, soil microbial biomass was little affected by the passage of soil through the gut of O. lacteum. The soil microflora of the field, the 5-, 12-, and about 50-year-old fallow was limited by carbon, whereas in the beechwood soil phosphorus limited microbial growth. NItrogen availability to the soil microflora was low in the two younger fallow sites and high in the field and the third fallow. In the beechwood soil nitrogen supply did not affect microbial carbon utilization. Application of phosphorus stimulated glucose mineralization in the soil of the field, the third fallow, and the beechwood, but not in the two younger fallow sites. Therefor, the nutrient status of the soil microflora seems to have changed during secondary succession: presumably, during the first phase the availability of nitrogen decreased, whereas during the second phase microbial phosphorus supply became more important, which resulted in phosphorus limitation of the soil microflora in the climax ecosystem. The passage of soil through the gut of O. lacteum caused an alteration in the microbial nutrient status. Generally, microbial growth in earthworm casts was limited by carbon. The relative effect of the gut passage of the soils on microbial carbon utilization seems to increase during succession. Therefore, the effect of decomposer invertebrates on microbial nutrient supply seems to increase during secondary succession. In general, nitrogen did not limit microbial carbon utilization in earthworm casts. Phosphorus requirements of the soil microflora were lowered by the gut passage of the soil of the third fallow site and the beechwood, which indicates an increased phosphorus supply in earthworm casts. Howerver, this additional supply was not sufficient to enable optimal carbon utilization by the soil microflora. The results indicate that the effect of decomposer invertebrates on the soil microflora depends on the nutrient status of the ecosystem.