DOI: 10.5061/dryad.8w9ghx3w9
title: Mesofauna and macrofauna densities at species/group level from 2008 to 2020 in three regions in Germany

description: Global biodiversity loss is threatening ecosystem functioning and human
 well-being. Arthropods above the ground have substantially decreased in
 abundance and diversity during the last 15-20 years. However, changes in
 belowground biodiversity, particularly in forests, received little
 attention. Here, we analysed a comprehensive dataset of soil-living meso-
 and macrofauna in forests differing in land-use intensity within the
 framework of the open research platform ‘Biodiversity Exploratories’.
 Abundance of soil animal species was analysed at three-year intervals,
 covering 12 years from 2008 to 2020. Neither density and species richness
 nor gamma diversity of both soil meso- and macrofauna declined, suggesting
 contrasting dynamics of biodiversity above and below the ground. Soil
 mesofauna density and diversity varied significantly between years within
 regions, with the variations being closely related to soil moisture of the
 previous winter months and during sampling. While the stability of
 mesofauna and of some macrofauna taxa was strongly correlated with
 asynchrony of species-fluctuations, and in part with effective diversity,
 overall neutral or positive variance ratios suggested that most species
 fluctuated synchronously. These synchronous fluctuations were likely due
 to variations in abiotic conditions such as soil moisture and presumably
 overprinted biotic drivers of stability. Stability was not directly
 related to forest management, although for mesofauna, it differed between
 forest types within regions. While documenting an astounding resilience of
 soil animals in temperate forests to the ongoing biodiversity decline, our
 findings help to better understand temporal patterns of soil fauna density
 and diversity and the drivers of their community stability.
description: Samples were taken in three regions in Germany (Swabian Alb,
 Schorfheide Chorin, and Hainich-Dün), spring (April to June) from 2008 to
 2020 in intervals of three years, resulting in 5 sampling points. Samples
 were taken from 5 m × 5 m subplots located within the 100 m × 100 m
 gridplots as described in Erdmann et al. (2012) and Klarner et al. (2014).
 Briefly, the four forest types were replicated four times in each region,
 resulting in 48 sampled forests. In each forest, one soil core of 5 cm Ø
 was taken for the extraction of mesofauna and one soil core of 20 cm Ø for
 the extraction of macrofauna. We separated the litter layer (variable
 thickness) and the top 5 cm of the soil underneath for extracting meso-
 and macrofauna by heat (Macfadyen 1961). Individuals from the two layers
 were pooled for statistical analyses. Lumbricidae were sampled from an
 area of 0.25 m² from each forest using a combination of hand sorting and
 extraction with mustard solution. First, the litter layer was removed and
 checked manually for earthworm specimens. Then, a mustard solution
 consisting of 100 g mustard powder (Semen Sinapis plv.; CAELO, Cesar
 & Loretz GmBH, 40721 Hilden, Germany) dissolved in 10 l water was
 applied to the soil in 2 steps with initially 5 liters and another 5
 liters after 15 min (Eisenhauer <em>et al.</em> 2008).
 Emerging Lumbricidae were hand-collected for a total of 30 min. All
 animals were stored in 70% ethanol until identification. Mesofauna
 (Oribatida, Collembola, and Mesostigmata) and macrofauna taxa (Isopoda,
 Diplopoda, Lumbricidae, and Chilopoda) were identified to species level
 using appropriate keys (Eason 1964; Hopkin 2007; Karg 1989; Karg 1993;
 Krantz & Ainscough 1990; Oliver & Meechan 1993; Schubart
 1934; Sims & Gerard 1985; Weigmann 2006). Araneae and Coleoptera
 were only sorted at the taxon level and were not considered in the
 analyses of diversity. Precipitation for each year and
 sampling site was derived from the RADOLAN (Radar Online Adjustment)
 product of the German Weather Service (Deutscher Wetterdienst), which
 provides hourly radar-based precipitation estimates for Germany adjusted
 to rain gauge data on a 1 km² grid (Kreklow <em>et al.</em>
 2019). Temperature was measured with environmental sensors installed at 2
 m above the ground at all sites (Fischer <em>et al.</em>
 2010). We calculated mean values for temperature and precipitation in
 winter (December to February) and spring (March to May). We also used the
 silvicultural management intensity indicator (SMI), including a risk and
 density component, and accounting for tree species, tree age, and
 aboveground living biomass (Schall & Ammer, 2013; 2023); it was
 taken from the Biodiversity Exploratories database (BExIS). Microbial
 biomass in leaf litter and soil was assessed by measuring the maximum
 initial respiratory response (MIRR; mg O<sub>2</sub> g^−1
 h^−1) after glucose addition (SIR method; Anderson & Domsch 1978)
 in an automated O<sub>2</sub> micro-compensation apparatus
 (Scheu 1992). Glucose (80 and 10 mg g^−1 dry weight for litter and soil,
 respectively) was added as an aqueous solution to approximately 1 g of
 leaf litter material (Beck <em>et al.</em> 1997).
description: # Mesofauna and macrofauna densities at species/group level from 2008 to
 2020 in three regions in Germany
 [https://doi.org/10.5061/dryad.8w9ghx3w9](https://doi.org/10.5061/dryad.8w9ghx3w9) The data files are abundances of soil animals per sample (in case of mesofauna, i.e., Collembola, Oribatida, and Mesostigmata, taken from a 5 cm soil core, in case of macrofauna, i.e., Araneae, Chilopoda, Coleoptera, Diplopoda, Isopoda, Lumbricidae, taken from a 20 cm soil core, each including leaf litter and upper 5 cm of soil). In addition, one data file (BE-environment_2008-2020.csv.) includes environmental variables such as temperature, precipitation, and land-use intensity. We calculated density, diversity (richness, effective diversity, gamma diversity), and stability of soil fauna in three-year intervals from 2008 to 2020 and modeled temporal trends and environmental drivers of soil fauna abundance, diversity, and stability. ## Description of the data and file structure **Note on Data File Format** Although the data files are saved with the `.csv` extension, the values are separated by **semi-colons (`;`)** rather than commas. Please ensure that your software or import settings are configured accordingly when opening these files. #### BE_Oribatida_2008-2020.csv Spreadsheet with an abundance of Oribatida species per plot and year (taken from a 5 cm soil core with leaf litter and the upper 5 cm of soil) Column information: plotyear: Sampling site and year of sampling Abundance: Rowsums of the species abundance data given in columns "C" to "DV". Columns "C" to "DV": Currently valid scientific names of oribatid mite species and their individual abundance for each site and sampling year. LandScape: The three sampling regions in Germany [Swabian Alb (Alb), Hainich Dn (Hai), Schorfheide Chorin (Sch)]. forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (B150), actively managed mature stands (approx. 70 years, B70), and actively managed young stands (30 years, B30) Plot: Sampling site year: Sampling year #### BE_Collembola_2008-2020.csv Spreadsheet with an abundance of Collembola species per plot and year (taken from a 5 cm soil core with leaf litter and the upper 5 cm of soil) Column information: year: Sampling year Plot_ID: Sampling site forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (B150), actively managed mature stands (approx. 70 years, B70), and actively managed young stands (30 years, B30) Expl: The three sampling regions in Germany [Swabian Alb (Alb), Hainich Dn (Hai), Schorfheide Chorin (Sch)]. sum: Rowsums of the species abundance data given in columns "F" to "DB". Columns "F" to "DB": Currently valid scientific names of collembolan species and their individual abundance for each site and sampling year. #### BE_Mesostigmata_2008-2020.csv Spreadsheet with an abundance of Mesostigmata species per plot and year (taken from a 5 cm soil core with leaf litter and the upper 5 cm of soil) Column information: EP_Plotid: Sampling site sampling_year: Sampling year expl: The three sampling regions in Germany [Swabian Alb (Alb), Hainich Dn (Hai), Schorfheide Chorin (Sch)]. forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (B150), actively managed mature stands (approx. 70 years, B70), and actively managed young stands (30 years, B30) code: Different coding for Coniferous forests (Fi/Ki),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (Bu nat), actively managed mature stands (approx. 70 years, Bu alt), and actively managed young stands (30 years, Bu jung) region: Different coding for the three sampling regions in Germany [Swabian Alb (AEW), Hainich Dn (HEW), Schorfheide Chorin (SEW)]. Columns "G to "DF": Abbreviations of scientific names of mesostigmatid mite species and their individual abundance for each site and sampling year. #### *BE_Macrofauna_2008to2020_wide.csv* Spreadsheet with an abundance of macrofauna (Isopoda, Diplopoda, Chilopoda, Lumbricidae) species per plot and year (taken from a 20 cm soil core with leaf litter and the upper 5 cm of soil) Column information: year: Sampling year Region: The three sampling regions in Germany [Swabian Alb (AEW), Hainich Dn (HEW), Schorfheide Chorin (SEW)]. EP_PlotID: Sampling site forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (B150), actively managed mature stands (approx. 70 years, B70), and actively managed young stands (30 years, B30) Columns "E to "CG": Abbreviations and scientific names of macrofauna species and their individual abundance for each site and sampling year. Note that earthworms (Lumbricidae) have non-integer numbers as abundances of juveniles from a genus were assigned to relative abundances of adult species in the same region/forest type/year. #### *BE_Chilopoda_specieslvl_2008-2020.csv* Spreadsheet with an abundance of Chilopoda species per plot and year (taken from a 20 cm soil core with leaf litter and the upper 5 cm of soil) Column information: year: Sampling year Region: The three sampling regions in Germany [Swabian Alb (AEW), Hainich Dn (HEW), Schorfheide Chorin (SEW)]. EP_PlotID: Sampling site forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (Bu nat), actively managed mature stands (approx. 70 years, Bu alt), and actively managed young stands (30 years, Bu jung) Columns "E to "AB": Scientific names of Chilopoda species/taxa and their individual abundance for each site and sampling year. #### *BE_Diplopoda_specieslvl_2008-2020.csv* Spreadsheet with an abundance of Diplopoda species per plot and year (taken from a 20 cm soil core with leaf litter and the upper 5 cm of soil) Column information: EP_PlotID: Sampling site year: Sampling year Region: The three sampling regions in Germany [Swabian Alb (AEW), Hainich Dn (HEW), Schorfheide Chorin (SEW)]. forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (Bu nat), actively managed mature stands (approx. 70 years, Bu alt), and actively managed young stands (30 years, Bu jung) Columns "E to "AC": Scientific names of Diplopoda species/taxa and their individual abundance for each site and sampling year. Class: Taxonomic group (i.e., Diplopoda) #### *BE_Isopoda_specieslvl_2008-2020.csv* Spreadsheet with an abundance of Isopoda species per plot and year (taken from a 20 cm soil core with leaf litter and the upper 5 cm of soil) Column information: EP_PlotID: Sampling site year: Sampling year Region: The three sampling regions in Germany [Swabian Alb (AEW), Hainich Dn (HEW), Schorfheide Chorin (SEW)]. forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (Bu nat), actively managed mature stands (approx. 70 years, Bu alt), and actively managed young stands (30 years, Bu jung) Columns "E to "O": Scientific names of Isopoda species and their individual abundance for each site and sampling year. Class: Taxonomic group (i.e., Isopoda) #### *BE_Lumbricidae_specieslvl_2008-2020.csv* Spreadsheet with an abundance of Lumbricidae species per plot and year (taken from a 20 cm soil core with leaf litter and the upper 5 cm of soil) Column information: EP_PlotID: Sampling site year: Sampling year Region: The three sampling regions in Germany [Swabian Alb (AEW), Hainich Dn (HEW), Schorfheide Chorin (SEW)]. forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (Bu nat), actively managed mature stands (approx. 70 years, Bu alt), and actively managed young stands (30 years, Bu jung) Columns "E to "R": Scientific names of Lumbricidae species and their individual abundance (rounded, see non-integer abundances in *BE_Macrofauna_2008to2020_wide*; abundances &lt;1 were rounded to 1, all other abundances were rounded to the closest integer) for each site and sampling year. Class: Taxonomic group (i.e., Annelida) #### *Macrofauna_grouplvl.csv* Spreadsheet with an abundance of macrofauna at the taxonomic group level (Araneae, Chilopoda, Coleptera including larvae, Diplopoda, Isopoda, Lumbricidae) per plot and year (taken from 0.25 m using mustard extraction) in long-format, including environmental variables. Column information: plotyear: Sampling site and year of sampling Plot_ID: Sampling site Group: taxonomic group region: The three sampling regions in Germany [Swabian Alb (AEW), Hainich Dn (HEW), Schorfheide Chorin (SEW)]. abundance_plotlvl: abundance per plot (taken from a 20 cm soil core with leaf litter and upper 5 cm of soil for all taxa except Lumbricidae, and from 0.25 m using mustard extraction for Lumbricidae) forest.type: Coniferous forests (Conif),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (Bu nat), actively managed mature stands (approx. 70 years, Bu alt), and actively managed young stands (30 years, Bu jung) year: Sampling year Precipitation_winter_mean: Mean values for precipitation (mm) in winter (December to February) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). airtemp_2m_above_winter_mean: Mean values for air temperature (°C) at 2 meters height in winter (December to February) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). Precipitation_wintersum: Mean values of the total precipitation (mm) in winter (December to February) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). airtemp_2m_above_spring_mean: Mean values for air temperature (°C) at 2 meters height in spring (March to May) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). Precipitation_spring_mean: Mean values for air temperature (°C) at 2 meters height in spring (March to May) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). Precipitation_spring_sum: Mean values of the total precipitation (mm) in spring (March to May) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). SMIr: Derived from a function of tree species and stand age. SMIr quantifies the age-dependent probability of losing a stand before it reaches 180 years. SMId: Derived from a function of the silvicultural regime, stand age, and tree species.SMId accounts for the deviation of the actual stocking from a fully stocked mature forest in terms of basal area, which has been modified by harvests and thinning. SMI: Combining SMIr and SMId (values range from 0 -1). Values approach 0 in unmanaged forests. Note that only SMI was used in the analysis. Data in columns ("SMIr", "SMId", "SMI") were sourced from: * Schall, P., Ammer, C. How to quantify forest management intensity in Central European forests. *Eur J Forest Res* **132**, 379396 (2013). [[https://doi.org/10.1007/s10342-013-0681-6\\](https://doi.org/10.1007/s10342-013-0681-6\\) ]([https://doi.org/10.1007/s10342-013-0681-6\\](https://doi.org/10.1007/s10342-013-0681-6\\) * Schall, Peter; Ammer, Christian (2023): SMI annual dynamics - Silvicultural Management Intensity Dynamics on all forest EPs, 2008 - 2020. Version 9. Biodiversity Exploratories Information System. Dataset. [https://www.bexis.uni-jena.de(opens in new window)](https://www.bexis.uni-jena.de/). Dataset ID= 31217 Cmic_litter: Microbial biomass per g of leaf litter (g Cmic g-1). Cmic_soil: Microbial biomass per g of soil (g Cmic g-1). pred_dec: trophic group, i.e., predator or decomposer abund_m2: abundance per m #### *BE_environment_2008-2020.csv* Spreadsheet with environmental factors per plot and year Column information: plotyear: Sampling site and year of sampling plotID1:Sampling site year: Sampling year Precipitation_winter_mean: Mean values for precipitation (mm) in winter (December to February) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). airtemp_2m_above_winter_mean: Mean values for air temperature (°C) at 2 meters height in winter (December to February) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). Precipitation_wintersum: Mean values of the total precipitation (mm) in winter (December to February) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). code: Different coding for Coniferous forests (Fi/Ki),near-natural beech forests, unmanaged for at least 60 years, with an approximate tree age of 150 years (Bu nat), actively managed mature stands (approx. 70 years, Bu alt), and actively managed young stands (30 years, Bu jung) region: The three sampling regions in Germany [Swabian Alb (AEW), Hainich Dn (HEW), Schorfheide Chorin (SEW)]. nr: Different coding for sampling site plotid2: Different coding for sampling site and year of sampling airtemp_2m_above_spring_mean: Mean values for air temperature (°C) at 2 meters height in spring (March to May) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). Precipitation_spring_mean: Mean values for air temperature (°C) at 2 meters height in spring (March to May) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). Precipitation_spring_sum: Mean values of the total precipitation (mm) in spring (March to May) calculated from hourly measurements of the RADOLAN (Radar Online Adjustment) product of the German Weather Service (Deutscher Wetterdienst). SMIr: Derived from a function of tree species and stand age. SMIr quantifies the age-dependent probability of losing a stand before it reaches 180 years. SMId: Derived from a function of the silvicultural regime, stand age, and tree species.SMId accounts for the deviation of the actual stocking from a fully stocked mature forest in terms of basal area, which has been modified by harvests and thinning. SMI: Combining SMIr and SMId (values range from 0 -1). Values approach 0 in unmanaged forests. Note that only SMI was used in the analysis. Data in columns ("SMIr", "SMId", "SMI") were sourced from: * Schall, P., Ammer, C. How to quantify forest management intensity in Central European forests. *Eur J Forest Res* **132**, 379396 (2013). [[https://doi.org/10.1007/s10342-013-0681-6\\](https://doi.org/10.1007/s10342-013-0681-6\\) ]([https://doi.org/10.1007/s10342-013-0681-6\\](https://doi.org/10.1007/s10342-013-0681-6\\) * Schall, Peter; Ammer, Christian (2023): SMI annual dynamics - Silvicultural Management Intensity Dynamics on all forest EPs, 2008 - 2020. Version 9. Biodiversity Exploratories Information System. Dataset. [https://www.bexis.uni-jena.de(opens in new window)](https://www.bexis.uni-jena.de/). Dataset ID= 31217 Cmic_litter: Microbial biomass per g of leaf litter (g Cmic g-1). Cmic_soil: Microbial biomass per g of soil (g Cmic g-1).