Development of soil fauna at mine sites during 46 years after ...

Pedobiologia 45, 243–271 (2001)© Urban & Fischer Verlaghttp://www.urbanfischer.de/journals/pedo

0031–4056/01/45/03–243 $ 15.00/0

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Wolfram Dunger and Manfred Wanner**with H. Hauser, K. Hohberg, H.-J. Schulz, T. Schwalbe, B. Seifert, J. Vogel, K. Voigtländer, B. Zimdars and K. P. Zulka1

State Museum of Natural History, POB 300 154, D-02806 Görlitz, Germany 1 Institute of Zoology, University of Vienna, Althanstr. 14, A-1090 Vienna, Austria

Submitted: 7. July 2000Accepted: 1. November 2000

Summary

At the afforested overburden heap „Langteichhalde“ (lignite mining district Berzdorf,Eastern Germany) a long-term study of the soil fauna (since 1961) was completed bynew investigations between 1996-1999 taking microfauna, mesofauna and macrofa-una into consideration. Detailed information about the present development of testateamoebae, nematodes, lumbricids, spiders and harvestmen, oribatids, centipedes, mil-lipedes, apterygotes, ants, carabid and staphylinid beetles are briefly compared withthe earlier succession of these groups. Generally, both the species inventory as well asdiversity points to a change from open landscape communities to woodland associati-ons, albeit in a more or less poor condition. Differences of the settlement dependingon deciduous or coniferous afforestation type and surface profile (crests, troughs) arediscussed.

The contribution of soil organisms to organic matter decomposition was studied bybait lamina tests, the minicontainer technique, and by calculating the potential zooticdecomposition level. At the deciduous afforestation the overall soil biological activityreaches the level of natural woodland soils very quickly (after 10 - 20 years), eventhough based upon the continuing development of soil faunal entities. Primarily coni-ferous afforestations show an impeded decomposing activity, even if developing in a

* Supported by the German Federal Ministry of Education and Research (BMBF, FKZ 0339668)**E-mail corresponding author: [email protected]

244 Wolfram Dunger et al.

mixed wood. During succession of the mine site community an interesting interrelati-onship between parts of the soil fauna, especially between lumbricids and microar-thropods, can be demonstrated.

Key words: Soil micro-, meso-, macrofauna, opencast mining dumps, primary suc-cession, decomposition, minicontainer, bait lamina test

Introduction

During the 20th century the increasing demand for energy has led worldwide, but es-pecially in Germany, to the opening of mining areas at dimensions of thousands ofkm2. One of the outcomes of this cataclysm is the increased number of summarisingbooks on the recultivation of post-mining landscapes at the end of the century (Hüttlet al. 1996, 1999; Pflug 1998; Broll et al. 2000). Concerning the succession of ani-mals, the first worldwide view was edited by Majer (1989).

In Germany, two main study centres have been established, the Lower Rhine mi-ning region (Glück 1989; Topp et al., in press) and the Lusatian mining district, divi-ded into Lower Lusatia with its centre at Cottbus (Hüttl et al. 1999) and Upper Lusa-tia with its centre at Görlitz (Dunger 1968; Dunger et al., in prep.). Stimulations to in-vestigations on the immigration and development of soil fauna at freshly deposited li-gnite mine spoils goes back to a discussion with Wilhelm Kühnelt at the University ofLeipzig in 1956. There were two reasons to take up such studies: From the viewpointof soil biological ecology the open cast coal mines offered a tremendous experimen-tal field to study soil animals in primary succession and, seen from the viewpoint ofrecultivation being very poorly developed at this time, knowledge of the soil fauna de-velopment was required as an indicator for artificial cultivation techniques. Later on,as a further point of interest, results from the soil faunal development on mine siteareas became important for a theoretical approach to primary succession (Dunger &Wanner 1999a).

To obtain results fulfilling these objectives, the State Museum of Natural Historyat Görlitz started a long-term programme for studying the soil fauna at mine sites asearly as in 1960. The objectives of this study, continued until 1999, were as follows:

– What are the specific characteristics of mine soils as habitats for soil animals andwhich of the peculiarities of the deposited substrates as well as of the rehabilitationtechniques are important for the soil fauna?

– Which kind of immigration type is used by the soil fauna and how much time isnecessary for the immigration of different taxa?

– Which successional direction is taken by the members of the soil fauna and howdevelop single populations and complex communities?

– What is the importance of different kinds of faunal activities in development anddifferentiation of soil ecosystems at mine soils?

– Which role plays the soil fauna in supporting a fast and efficient rehabilitation ofmined areas and what are the most adequate human techniques in recultivation to fo-ster this activity?

245Soil fauna at mine sites

The soil faunal studies of the Museum Görlitz started at mine sites at the Germansouth-eastern brown coal mining district of Berzdorf with comparative studies at themining areas of Lower Lusatia (Dunger 1979) and south of Leipzig (Brüning et al.1965; Dunger 1969). Within this project, a long-term study of the soil fauna of theoverburden heap „Langteichhalde“ (LTH) at the mining district of Berzdorf south ofGörlitz was established in 1960. Successive results of this project have been publis-hed by Dunger (1968, 1987, 1989, 1997a, b, 1998a, b). This report presents new in-formation from a study between 1996-1999 comprising not only continued investi-gations up to 46 years following the mine site recultivation but also an extension ofthe groups studied taking the microfauna (testate amoebae, nematods) into considera-tion for the first time.

Materials and Methods

Study sites

The study was carried out mainly at a dump („Langteichhalde“, LTH) of the brown coal opencast mining of Berzdorf (south of Görlitz), Upper Lusatia, Eastern Germany. At this dump(established between 1951 and 1955) the mine site A was afforested in 1952 with Alnus gluti-nosa, Populus sp. (hybr.) and Robinia pseudacacia, and mine site L with Pinus sylvestris. Forcomparative purposes, the following test sites were investigated: at the same dump the minesite H, afforested as site A in 1955, the mine site T at an adjacent dump afforested as A in 1959and the youngest mine site N at a dump 7 km north of site A, afforested with Populus sp. in1961. Further afforested young mine sites were studied 1997/99 4 km to the west of site A („In-nenkippe“, IK).

The soil cover on the final dump consists mainly of sands of Pleistocene and Tertiary ori-gin interspersed with lignite sands and dark Tertiary loam and clay. Amelioration was mainlyby liming, the pH(KCl) in the upper 10 cm was 5.8 in 1963 decreasing to 5.2 in 1998. Some 15years after afforestation in mine site A a first woody stage with final soil shading was reached.Results of top soil investigations in 1998 are for site A: humus form L-mull, ectohumus cover1 cm, Ah 0-10 cm; for site L: humus form moder, ectohumus cover 3 - 7 cm, Ah 0-2 cm; detai-led soil characteristics are shown in Table 1.

Table 1. Soil properties of the Berzdorf mine sites A (deciduous afforestation) and L(pinus afforestation tending to a mixed wood). Data for 1963 from Dunger (1968);data for 1998 from Hauser & Kowarsch (1998) and Wanner & Dunger (1999). c=crest, t= trough; x= mean value; a= 0-5 cm, b= 5-10 cm for the 1998 data; a= 0-2 cm,b= 5-7 cm for the 1963 data. SOM= soil organic matter incl. lignite (%), Vp= pore vo-lume (%). The other data from 1998 derived from 0-10 cm soil depth

Site date pH(KCl) Ct N S C/N SOM Vpca cb ta tb c t x c t x

A 1963 5.8 5.9 5.5 4.9 10.6 5.6 8.1 44.1 47.9 46.01998 5.1 5.1 4.3 4.2 5.0 0.2 0.001 20.0 9.9 48.4

L 1998 5.1 4.1 4.4 4.0 2.3 0.1 0.007 22.0 4.5 49.8


Methods

Protists and nematodes were sampled in spring and autumn 1997 and 1998 resp. by soil cores(Ø 5cm, 0-5 and 5-10 cm depth, 5 replicates) and examined microscopically using an invertedmicroscope. Testate amoebae were examined directly (Wanner & Dunger 1999); nematodeswere extracted using a modified Baermann-funnel.

Edaphic microarthropods („Berlese-fauna“) were sampled at 6 dates altogether in au-tumn 1997 and spring - autumn 1998 with 50 soil cores (Ø3.5cm) at each of mine sites A andL, divided into 0-5 and 5-10 cm depth and soil profile. There were 12 replicates for crests,and 13 replicates for troughs; the fifth and sixth sampling dates in autumn 1998 are based onpooled data-sets, thus the first four samplings were available for statistical analysis. The ani-mals were extracted by thermoeclectors of a modified Tullgren-type (Dunger & Fiedler(1997).

Epedaphic micro- and macroarthropods were collected by pitfall traps (each trap was ana-lysed separately) at 6 dates from autumn 1997 up to autumn 1998 with 9 traps at each of minesites A and L, divided into soil profiles crests (four traps) and troughs (five traps). These data-sets were used for statistical analysis. Additional animals came from area samples and soilmacro-cores (see earthworms).

Earthworms were calculated on the basis of 60 area samples (0.25 m2, three replicates onthe troughs, two replicates on the crests; arithmetic means were used for m2-calculations) usingdiluted formalin in combination with 80 soil macro-cores sorted by hand (Dunger & Fiedler1997) at all of mine sites A and L (five replicates on crests and on troughs, Ø 9.5 cm).

Decomposition was assessed by two methods: The bait lamina test was used (after Törne1997) with a bait substance of cellulose (65%), agar agar (15%) and wheat bran (10%). Fourtest sites (A and L; crest and trough, resp.) with three plots of 16 bait laminas were established.Parallel to the bait lamina, the minicontainer test (Eisenbeis 1998) was carried out by inserting10 rods each in crests and troughs at each of the mine sites A and L from October 1997 to Ja-nuary 1999. At 6-weekly intervals at each stand one rod, containing 4 minicontainers with2000, 500 and 20 µm mesh size resp., were removed, the inhabitants extracted by a ther-moeclector, and the remaining organic material (poplar litter) weighed. Thus a period of 60weeks was investigated. Because of the preliminary character of the minicontainer test (lowamount of replicates), a detailed statistical analysis was omitted.

Litter production was tested by three litter samplers (1 m2 area) at each site A and L bet-ween May 1998 and May 1999 being emptied 10 times. For weighing, the litter was dried at60°C. Additionally the field layer was harvested at 3 plots of each site A and L in May and Sep-tember 1998.

Metabolic equivalent values of lumbricids (MELu) and microarthropods (MEMa), and therespective potential zootic decomposition level (DLZpot) were calculated according to Dunger& Fiedler (1997):

ME = B‘ * k * 102 * Y/X where B‘ is the mean biomass (g wet mass m-2), k is the caloric equivalent factor (20*103 J ml-1 O2), X is the mean living mass per individual (g), and Y is the oxygen consumption(laboratory data, ml O2 ind-1 h-1 at 10°C).

From this the potential zootic decomposition level (DLZpot) can be deduced thus:DLZpot=DZpot*100/H

where DZpot is the total of ME of an animal group and H is the yearly supply of organic mattermeasured as annual litter production incl. field layer harvest (KJ m-2 a-1).

Analogously, respiratory equivalences (RE) are calculated according to Dunger (1991b):RE=B‘ * 104 * Y/X.Data were evaluated with an analysis of variance (ANOVA; normality was tested with the

Kolmogorov-Smirnov test). If input data (as mentioned above) revealed heterogeneous varian-ces (even after transformation) only those factors with P<0.01 were considered, or nonpara-


metric tests were used (Mann-Whitney test and Kruskal-Wallis one-way analysis of variance).The requirements for statistics were met according to Bauer (1986) and Sachs (1999). The soft-ware package SPSS was used for all statistics.

Results

Components of the soil fauna – testate amoebae

The data presented are based on soil samples (0-5 cm and partly 5-10 cm depth), eit-her pooled (5 soil cores per sampling date, from 1996-1998) to reveal seasonal andsuccessional trends, or counted separately from three or five replicates to provide in-formation about variability within a study site. Generally, small, rapidly reproducingubiquists (r-strategists) predominated the afforestations (with the exception of Difflu-gia stoutii Ogden – as far as we know the first record for Germany and the second re-cord since the original description in 1983). During the initial stage, testate amoebaecolonised the substrate „additively“, i.e. without replacing other testate amoebae taxa,within a few months. Regarding two up to 46-year-old afforested mine soils (IK, LTH:especially the Pinus sites), a consistent development was observed in the amoebalspecies inventory (all mine sites: 48 taxa; LTH-L: 35; LTH-A: 36 species), populationdensities and biomasses in relation to age, substrate and stocking of the afforestations.Abundances and biomasses were of the same order, or even higher, as those describedfor undisturbed forest soils (e.g. compiled in Foissner 1987). Six out of 48 taxa con-tributed 61 to 87 % to the mean total amoebal density (26-366 x106 ind. m-2; 0.4-4.8 gm-2; mean values). Species richness increases, as a rule, in the older test sites (Wanner& Dunger 1999, 2001). However, typical humus-inhabiting, large-sized testate amo-ebae (e.g. Trigonopyxis arcula (Leidy), Hyalosphenia spp., Nebela spp.) were lackingor occurred rarely. With respect to afforestation, abundances and biomasses were re-markably higher in the Pinus afforestations (100-238 x106 ind. m-2; 0.7-3.0 g m-2) ascompared to the deciduous sites (47-89 x106 ind. m-2; 0.4-0.9 g m-2; mean values).Species richness, abundances and biomasses on LTH-L (Pinus site) were significantlyhigher as compared to LTH-A on the second sampling date (deciduous site, P<0.01,Oct. 1998), while the soil surface (crests and troughs) had no influence (P>0.05; non-parametric H-test, for both sampling dates). Meisterfeld (1997) found, as shown inthis study, no „typical“ humus-inhabiting species of testate amoebae on reclaimedmine soils in the lignite district of the Rhineland. Only more or less ubiquitous, smallspecies became established. Balík (1996) investigated testate amoebae in the Sokolovcoal mine district and described, similarly to our data, a rapid development of the te-state amoebae assemblages in the initial stage. Different ages and stages of recultiva-tion are characterised by distinct soil testate amoebae assemblages (Wanner et al.1998, 1999; Wanner & Dunger 1999, 2001).

Components of the soil fauna – nematodes

The investigation (for details see Hohberg, in prep.) is based on soil samples (0-5 cmand partly 5-10 cm depth), taken in five replicates in spring and autumn 1998. Gene-rally, nematode density and biomass of afforested Berzdorf mine sites (IK, LTH) werelow (0.06 – 1.22 x 106 ind. m-2; 1.9 – 78.1 mg m-2 ), however still within range of


literature data from natural temperate forest soils. Considering the relatively high den-sities of testate amoebae mentioned above, it seems improbable that a possible lownutrient supply of mine site soils might cause the rather low nematode densities. Weassume that detrimental effects of soil structure, vegetation cover and specific preda-tors (e.g. the tardigrade Macrobiotus richtersi Murray; see Hohberg, in prep.) shouldbe taken into consideration. Altogether, in nine mine soils investigated in spring 1998,119 species were recorded, belonging to 55 genera. With 56 species (deciduous site A)and 35 species (primarily coniferous site L), species numbers of the 46-year-old affo-restations were comparable to those recorded from natural forest soils. Alder affore-stations (IK) especially, provided favourable conditions for the coexistence of manyspecies, whereas coniferous as well as poplar plantations resulted in lower nematodediversity. Bacterivores dominated (above all Acrobeloides nanus (De Man) on pineand poplar sites; Plectus spp. and Panagrolaimus spp. on alder sites and deciduoussite A). Fungi-/radicivorous species showed higher proportions in site L, whereas ob-ligate herbivorous species were more frequent in site A. Regarding two up to 46-year-old afforested mine soils (especially the pine series) the species inventory (mainlybacterivorous species) and the proportion of bacterivores increased. Neither maturityindex nor colonizer persister classes provided a useful bioindication for the long termsuccession of the Berzdorf mine site ecogenesis.

Components of the soil fauna – lumbricids

Lumbricid data are based on 60 „formalin samples“ and 80 hand-sorted soil samples(LTH-A, L) taken between autumn 1997 and 1998. The mean population densities andbiomasses were in LTH-A 524 ± 193 ind. m-2 / 83.1 ± 38.9 g(wm) m-2 and in LTH-L218 ± 143 ind. m-2 / 52.7 ± 50.0 g(wm) m-2. Epigeic species were Dendrobaena octa-edra (Savigny), which had been recorded since 1961, Dendrodrilus rubidus rubidus(Savigny), since 1985, and Lumbricus rubellus rubellus Hoffmeister, since 1961.Dendrodrilus rubidus was distributed extremely heterogeneously, and adult individu-als were totally absent in spring 1998 („L“) and autumn 1998 („A“). More importantwas L. rubellus, which surpassed 4-6 times („L“) up to 15 times („A“) the biomass ofthe „Dendrobaena-complex“. Generally, epigeic species shared 37% of the total lum-bricid density and 25% of the biomass in „A“ and 61% (ind.) 34% (biomass) in „L“,respectively (Table 2a). Lumbricus terrestris L. is the only anecic species which hasoccurred at site A since 1966. With respect to its ecological significance, as much as36% („A“) and 45% („L“) of the total lumbricid biomass and 18% („A“) and 10%(„L“) of the total abundances could be ascribed to L. terrestris in 1998. Aporrectodealonga longa Ude was detected only in 1966. Endogeic species have been found since1961 (Aporrectodea caliginosa caliginosa (Savigny), Aporrectodea rosea rosea (Sa-vigny), and Octolasion tyrtaeum (Savigny)). Two species occurred only occasionally:Allolobophora chlorotica chlorotica (Savigny) in 1966 and Octolasion cyaneum (Sa-vigny) in 1985. Adult A. caliginosa reached higher biomasses than A. rosea, althoughmost individuals of this genus are found in the juvenile stage. All endogeic species re-ached 40% („A“) and 21% („L“) of the total lumbricid biomass and 45% („A“) and27% („L“) of the total lumbricid density in 1998 (Table 2a). Total lumbricid densitiesand biomasses (yielded by formalin extraction) differed significantly between vegeta-tion (note the significant contribution of the endogeic life-forms), but not between soilsurface relief (Table 2b). It should be taken into consideration that differences in the


soil surface (which decreased gradually after dumping) remained more pronounced inthe A site. Furthermore, due to considerable variations in densities and biomasses, e.g.in 1997-1998, detailed estimations are difficult, and heterogeneity of the soil substrateproduce marked variations in population structure. However, during the first decadesof succession, troughs had been clearly preferred by earthworms, especially epigeiclife-forms. This (statistically non-significant) trend is still visible in 1999 (Table 2a,b). The observed species inventory (7 persistent species) is typical for disturbed areasand similar to „decomposer communities“ (Graefe 1993). Unsuccessful colonisationattempts of three other species (A. longa, A. chlorotica, O. cyaneum) seem to demon-strate that the ecological capacity is exhausted for the given period.

Components of the soil fauna – spiders and harvestmen

Pitfall trapping, hand sorting and soil core extraction (see methods) produced 51 spe-cies of Araneae and 9 species of Opiliones. Activity density (individuals per trap andweek) was highest in troughs of site A, especially with regard to harvestmen (Table3a, b). The arachnid assemblages of the 46-year-old sites (LTH-L, A) comprised morespecies as compared to the younger afforestations and differed in species composition.

Table 2a. Lumbricids: spectrum of life-forms, mean densities and biomasses (Ind.,BM (g wm) m-2). Six sampling dates (Oct. 1997-Oct. 1998), data combined from for-malin extraction and hand-sorting. 46-year-old Berzdorf mine sites (LTH-A: deci-duous forest; LTH-L: primarily afforested with pine) with crest- and trough-structu-red soil relief

LTH-A LTH-Lcrest trough crest trough

Ind. BM Ind. BM Ind. BM Ind. BM

epigeic earthwormsLumbricus rubellus, adult 18 8.3 19 6.3 3 1.9 8 4.2Lumbricus rubellus, total 87 18.0 121 19.3 12 12.1 30 17.6Dendrobaena s.l., total 67 1.2 113 2.3 86 2.1 133 4.3total epigeic 154 19.2 234 21.9 98 14.2 163 21.9anecic earthwormsLumbricus terrestris, adult 5 20.8 7 15.9 3 9.5 5 14.1total anecic 74 30.6 110 29.3 13 19.7 26 27.6endogeic earthwormsAporrectodea spp., total 136 23.3 157 24.1 52 10.1 43 10.0Octolasion tyrtaeum 92 9.5 90 8.3 16 0.7 7 1.1total endogeic 228 32.8 247 32.4 68 10.8 50 11.1

total 456 82.6 591 83.6 179 44.7 239 60.9

share (%) to the totalepigeic lumbricids 33.8 23.2 39.6 26.2 54.7 31.8 68.2 36.1anecic lumbricids 16.2 37.0 18.6 35.0 7.3 44.1 10.9 45.5endogeic lumbricids 50.0 39.7 41.8 38.8 36.0 24.2 20.9 18.3


Also, the proportion of woodland species (e. g. Bathyphantes nigrinus (Westring), Co-elotes inermis (L. Koch)) was higher, those of open landscape species lower, andpioneer species were no longer present. The overall species diversity indicates a rela-tively poor arachnid community structure as compared to naturally grown woodland.Moreover, the conspicuous presence of several woodland ecotone species (e.g. Tro-chosa terricola Thorell, cf. Heublein 1983) points to a more or less disturbed wood-land species assemblage of the oldest mine site afforestations. Generally, no distinctarachnid species assemblages were visible with regard to site or soil relief (trough orcrest). However, some single species apparently preferred humid troughs over thecrests (e.g. Gongylidium rufipes (L.), Pirata hygrophilus Thorell), while others, likeDiplocephalus picinus (Blackwell), almost exclusively occurred in the deciduouswoodland site A.

Components of the soil fauna – oribatids

Oribatid mites were examined by soil core extraction and, to a lesser extent, by pitfalltrappings (see methods). Mean population density at site A (26.5*103 ind. m-2 in 1985;24.4*103 ind. m-2 in 1997-1998) remained relatively constant in time, while at site L

Table 2b. Lumbricids: Statistical analysis based on data derived from formalin ex-traction only. “Total” means epigeic, anecic and endogeic species plus indetermina-ble juveniles from Lumbricus. Differences between surface (crests and troughs) or si-tes (A, L) are indicated by the level of significance of the nonparametric Kruskal-Wal-lis H-test

total epigeic anecic endogeic

site Ind 0.000*** 0.929 0.069 0.000***BM 0.000*** 0.464 0.045* 0.000***

surface Ind. 0.507 0.073 0.568 0.786BM 0.763 0.242 0.548 0.886

Table 3a. Arachnids from the Berzdorf mine sites LTH-A, L (see Table1); c=crests;t=troughs

Site (LTH) Ac At A Lc Lt L

Araneae (ind. trap-1week-1) 2.02 4.55 3.43 4.08 3.97 4.02Species trapped 21 22 27 23Add. species by hand sorting 2 5 4 4total 23 27 31 27

Opiliones (ind. trap-1week-1) 7.08 12.20 9.93 4.23 4.43 4.34Species trapped 6 6 7 5Add. species by hand sorting 1total 6 7 7 5


average density decreased (95.4*103 ind. m-2 in 1985; 18.3*103 ind. m-2 in 1997-1998;Table 4a). Oribatid mites, with a total of 90 taxa, indicated a high species diversity. 43species were common to both sides, while – in low abundance – 21 occurred exclusi-vely on site A and 26 species exclusively on site L. Species inventory, grouped afterBeck et al. (1997), points to a relatively cool-humid habitat type for the 46-year oldafforestations (Table 4b). The oribatid fauna depends strongly on vegetation and litterquality. In 1962, the population density at the Pinus site L exceeded by 16 times thedeciduous site A, and still 3.5 times in 1985. Owing to the increasing proportion of de-ciduous trees and the decreasing needle litter layer at site L, no striking differenceswere visible in 1997/1998. Species inventory on the deciduous site A pointed to amore or less open mixed wood habitat type (e.g. Ceratozetes gracilis (Michael), Me-dioppia obsoleta (Paoli), Galumna lanceata Oudemans), while species exclusivelyfound on the primary Pinus site L are not typical for coniferous habitat types.

With respect to soil surface properties, oribatid density was always (but partly non-significantly) higher on the crests as compared to the troughs (Table 4a; nonparame-tric H-test: pitfall traps: P=0.266 (soil surface) and P=0.123 (vegetation); Berlesedata: P=0.012** (soil surface) and P=0.931 (vegetation)).

Table 3b. Arachnids from the Berzdorf mine sites LTH-A, L (see Table3a). Results of the two-factorial ANOVA (data transformed) on the effects of „date” (factor 1, six samplings dates) and „surface” (factor 2, crests and troughs). Five replicates on the troughs, four replicates on the crests per site and date, n= 804/1541 (Aran./Opil.)

factor df MS F Sign. of F

Araneae (pitfall trappings)site LTH-Awithin cells 42 0.22„date” 5 4.32 19.76 0.000„surface” 1 2.97 13.58 0.001interaction 5 0.55 2.53 0.044site LTH-Lwithin cells 42 0.25„date” 5 5.61 22.83 0.000„surface” 1 0.006 0.02 0.879interaction 5 0.08 0.34 0.888

Opiliones (pitfall trappings)site LTH-Awithin cells 42 1.95„date” 5 9.67 4.96 0.001„surface” 1 17.41 8.93 0.005interaction 5 4.96 2.54 0.042site LTH-Lwithin cells 42 1.11„date” 5 3.21 2.89 0.025„surface” 1 0.00 0.00 0.961interaction 5 1.99 1.79 0.136


Components of the soil fauna – centipedes

Centipedes (11 species ) were studied by pitfall-traps, soil core extraction, sorting byhand, and minicontainers. At the deciduous site A, activity-density (0.81 ind. trap-1

week-1) was significantly higher than at site L (0.19 ind. trap-1 week-1; nonparametricH-test: P=0.000***). Population densities and biomasses indicated non-significantsite differences („A“ with 472 ind. m-2; 0.85 g (wm) m-2 and „L“ with 355 ind. m-2;0.99 g (wm) m-2; hand-sorting; nonparametric H-test (ind.): P=0.062). In contrast toearlier successional stages, a consistent influence of the surface profile (crests,troughs) on population density was not detectable in 1997/1998 ( hand-sorted mate-rial; H-test: P=0.215 (site L); P=0.614 (site A)). With regard to species inventory, Li-thobius microps Meinert, 1868 predominated in pitfall trappings, followed by the lessfrequent Lithobius forficatus (L.). The pioneer species Lamyctes fulvicornis Meinertwas completely lacking after 1965 on both mine sites. Four out of 11 species were tobe found exclusively at site A, of which three are geophilomorphs (Geophilus ins-culptus Attems, G. electricus (L.), Strigamia acuminata (Leach)). These subterraneancentipedes invade mine sites only after the formation of an at least minimal humic A-horizon. Furthermore, the first occurrence of Lithobius mutabilis C. L. Koch (seeDunger & Voigtländer 1990) indicates a progressive development from an open land-scape to a woodland community.

Table 4a. Oribatid mites, mean densities per soil core (9.08 cm2) and activity-densi-ties derived from pitfall-trapping, Berzdorf mine sites LTH-A, L (see Tables 1, 2)


Oribatids, total 28.60 17.61 18.92 14.54Oribatids, size class I (<0.5 mm) 25.55 16.28 23.25 18.34Oribatids, size class II (<1 mm) 1.25 1.18 1.44 0.42mean density (m-2) 29516 19185 24351 20837 11476 18282activity-density (ind. trap-1week-1) 1.81 1.87 1.84 7.54 1.35 4.10

Table 4b. Oribatid mites, species inventory according to higher taxonomic categories, Berzdorf mine sites LTH-A, L (see Tables 1, 2)

Site (LTH) LTH-A LTH-Lspecies % species %

Basic low oribatids 12 20.0 12 28.0Peripheral low oribatids 11 18.4 8 36.3Basic higher oribatids 3 0.2 1 0.2Eupheredermata 5 2.7 7 3.0Oppioidea 16 15.4 18 16.1Basic Pterogasterina 3 10.4 5 2.1Total oribatids 63 100 65 100


Components of the soil fauna – millipedes

Millipedes (9 species ) were studied using pitfall-traps, soil core extraction, sorting byhand, and minicontainers. On the deciduous site A, activity-density (2.68 ind. trap-1

week-1) was significantly higher than at site L (0.55 ind. trap-1 week-1; H-test:P=0.000***). At both sites, a remarkably increased activity-density has been obser-ved since 1961/1962 („A“: 0.44 ind. trap-1 week-1; „L“: 0.16 ind. trap-1 week-1). Po-pulation densities and biomasses now indicate inconsistent site differences („A“with201 ind. m-2; 1.31 g (wm) m-2 and „L“ with 110 ind. m-2; 3.06 g (wm) m-2; hand-sor-ting; H-test (ind.): P=0.066). In 1962, millipedes could hardly be detected by hand-sorting. Species inventory development has been documented by Dunger & Voigtlän-der (1990) from 1961. Since 1985/1986, species diversity decreased at site A and re-mained constant at site L; both sites now display a „deciduous forest-population“ do-minated by Glomeris hexasticha Brandt. Other species have been trapped indecreasing density: Julus scandinavius Latzel, Melogona voigtii (Verhoeff), Polyzo-nium germanicum Brandt, and Unciger foetidus (C. L. Koch). The pioneer speciesCraspedosoma rawlinsi Leach and Polydesmus inconstans Latzel had been nearlydisappeared at this stage. Generally, afforestation by deciduous woodland resulted ina typical diplopod community („A“), whereas primarily coniferous afforestation fol-lowed by a succession to a mixed forest caused a relatively species-poor community.The surface profile differentiation in crests and troughs has clearly decreased (especi-ally at site L) since dumping. Thus the observed trough preference in diplopod settle-ment during the first decades of succession has been diminished with time. In1997/98, diplopod activity-density in troughs remained higher only at site A (H-test:P=0.0003***; for site L: P=0.2963), and no differences were shown by hand-sortedmaterial (H-test: P=0.936 (site L); P=0.576 (site A)).

Components of the soil fauna – apterygotes

Springtails (51 species) and proturans (quantitative data only) were sampled by soilcore extraction and pitfall-trapping (collembolans only). With respect to collembolanpopulation density, no site or soil surface differences were visible (1997/1998; site A:28.3*103 ind. m-2; site L: 26.2*103 ind. m-2 (Table 5a, b); the smallest size class pre-dominates. In comparison to the 1985 samples, densities at site A remained constant(27.8*103 ind. m-2), while at site L they decreased slightly (35.9* 103 ind. m-2). Withrespect to pitfall-trapped epedaphic collembolans, activity in May 1997/1998 wasroughly as high as in 1962 or 1985, but decreased more distinctly in autumn. Consi-dering the 1997/1998 data, springtail activity-density displayed, in contrast to mostother tested groups, no significant site differences („L“, „A“; Table 5a, b). Proturandensity had decreased since 1985 on both study sites (1985/ 1997-1998: „A“: 2533/666 ind. m-2; „L“: 2670/ 732 ind. m-2). Collembolan species inventory revealed re-markable successional changes. Some edaphic springtails remained eudominant (Pa-risotoma notabilis (Schäffer), Mesaphorura macrochaeta Rusek, M. hylophila Rusek,and M. tenuisensillata Rusek), but formerly recedent species now became predomi-nant (e.g. Isotomiella minor (Schäffer), Megalothorax minimus Willem, Protaphor-ura armata (Tullberg), Stenaphorurella quadrispina (Börner), or Folsomia mano-lachei Bagnall). Characteristic species of typical associations for younger successio-nal stages decreased considerably (e.g. Proisotoma minuta (Tullberg), Micranuridapygmaea Börner, Isotomodes productus (Axelson)), or disappeared completely. As for


edaphic collembolans, epedaphic taxa have undergone considerable changes in spe-cies composition within the last 46 years. Dominant and characteristic species of ear-lier stages, such as Lepidocyrtus paradoxus Uzel, L. cyaneus Tullberg, Tomocerusvulgaris (Tullberg), Entomobrya multifasciata (Tullberg), Hypogastrura succineaGisin, and H. assimilis Krausbauer have been replaced by Tomocerus flavescens(Tullberg), Orchesella flavescens (Bourlet), and Sminthurinus aureus (Lubbock) in1997/1998. The observed changes in species composition indicate a replacement of anopen landscape community to that of a woodland. Typical sylvan species, such asNeonaphorura dungeri Schulz, Deuterosminthurus bicinctus (Koch), Dicyrtomafusca (Lubbock), or Arrhopalites sericus Gisin, appear now. Furthermore, differencesin species composition of sites L and A have been diminished during time in accor-dance with the development of „L“ to a mixed forest. With respect to collembolan po-pulation density, crests and troughs seemed to be colonised differently by epedaphiccollembolans, while no site differences for edaphic (0-5 cm soil cores) or epedaphicspringtails were visible in 1997/1998 (Table 5a, b). Furthermore, some species clearlypreferred deeper soil layers (e.g. Oligaphorura serratotuberculata (Stach)).

Components of the soil fauna – ants

Ant data are primarily based on nest records („A“: 29.9 nests 100 m-2; „L“: 24.5 nests100 m-2), and to a lesser extent on pitfall-trappings and soil cores. At site A, seven spe-cies were found, at site L only four. As first observed in 1985, the findings of1997/1998 confirm that open landscape-preferring pioneer species (Formica cinereaMayr, Lasius niger (L.), Tetramorium impurum (Förster)) are replaced by sylvan spe-cies. The recent woodland community is typical but poor in species, especially withregard to site L, where Lepidothorax spp. (normally common in Pinus forests) are to-tally lacking. This might be caused by the extraordinary dense field layer, keeping thesoil temperature at a low level. Species that are found by trapping only (Formica ci-nerea, Lasius niger) are assumed to be invaders from adjacent areas, while Myrmica

Table 5a. Apterygotes, mean densities per soil core (9.08 cm2) and m2; and activity-densities derived from pitfall- trapping, Berzdorf mine sites LTH-A, L (see Tables 1,2)


edaphic Collembola, total (soil core-1) 24.12 27.29 22.88 24.66

size class I (<0.5mm) 21.26 23.68 20.35 22.18size class II (<1 mm) 2.86 3.57 2.50 2.39size class III (< 2mm) – 0.04 0.03 0.09

edaphic Collembola, m-2 28309 26178epedaphic Collembola 14.0 17.4 15.9 9.7 18.1 14.3

(ind. trap-1week-1)

Protura, total (size class I only) 0.47 0.74 0.96 0.37

Protura, m2 666 732


rubra (L.), M. ruginodis Nylander (predominating at site „L“), Lasius platythoraxSeifert, L. flavus (Fabricius), and Stenamma debile (Förster) are also found in nests.Generally, ants are known to depend principally on epigeic habitat structures, and donot indicate the developmental stage of the soil.

Components of the soil fauna – carabid and staphylinid beetles

Ground beetles (site A: 21, site L: 19 species) and rove beetles (site A: 37, site L: 41species) were examined using pitfall traps and hand-sorting. Activity-density (indivi-duals per trap and week) of carabids (site A: 4.45, site L: 2.68) and staphylinids (siteA: 4.37, site L: 2.48) was always highest at the deciduous afforestation „A“ (H-test,P=0.009** (Carabidae, ind.) and P=0.002** (Staphylinidae, ind.)). In comparison tothe investigation period of 1985 (Vogel & Dunger 1991), at both sites the eurytopicwood-inhabiting ground-beetles Carabus nemoralis Müll. and Pterostichus oblongo-punctatus (F.) decrease, and the hygrophilic Leistus terminatus (Hellwig) appears forthe first time, becoming subdominant. Additional conspicuous replacements of spe-cies point to a change from an open landscape community to a woodland association.In accordance to the change of site L from a Pinus afforestation to a mixed forest, spe-cies inventory adjusted to site A, but some characteristic dominant species remainedat both sites (site A: e.g. Carabus hortensis L.; site L: Ocalea badia Er., Staphylinuserythropterus L.).

Contribution of soil organisms to decomposition of organic matter

The most important developmental process in mine site soil ecosystems is the forma-tion and the decomposition of organic matter. In this context, the vegetation develop-ment of the Berzdorf mining district is referred to by Dunger (1968); Dunger & Wan-ner (1999b) and Dunger et al. (in prep.).

According to the vegetation growth, the yearly amount of dead organic matterfluctuates during mine site succession. That can be shown by the evaluation of theabove soil primary net production (except that of the woody standing crop) measured

Table 5b. Collembola, mean densities per soil core (9.08 cm2) and m2; and activity-densities derived from pitfall- trapping, Berzdorf mine sites LTH-A, L. Results of thenonparametric ANOVA (Kruskal-Wallis H-test) on the effects of „surface” (crests andtroughs) and „site” (LTH-A, L). Soil cores: four sampling dates with 12/13 replicatesper crest/trough. Pitfall trappings: six sampling dates with 4/5 replicates percrest/trough

Epedaphic Collembola (pitfall trapping)„surface” (site A) P= 0.924„surface” (site L) P= 0.197„site” P= 0.890Euedaphic Collembola (soil core extraction)

0-5 cm 5-10 cm„surface” (site A) P= 0.544 P= 0.741„surface” (site L) P= 0.349 P= 0.037*„site” P= 0.900 P= 0.003**


as yearly litter production and field layer crop (Table 8). In 1998, the woody share oflitter was 34.7% at mine site A and 25.0% at mine site L; in the latter the needle lit-ter amounts to 56.7% of the total.

As an integrated approach to study litter decomposition the bait lamina test was in-troduced by Törne 1990 (Larink & Kratz 1994; Dunger & Fiedler 1997). It indicatesprimarily the biological activity in the litter layer and the upper soil layers, mainly in-volving both the participation of soil microorganisms as well as soil invertebrates. Thebait lamina test informs about relative overall biological activities within and bet-ween distinct test plots. In 1997 - 1998, bait lamina strips were exposed three times atfour test sites: crests (Ac) and troughs (At) at deciduous mine site A and crests (Lc)and troughs (Lt) at mixed woody mine site L. The results (Table 6) showed that theoverall biological activity was greater at troughs than at crests, especially at the Lmine site. Otherwise, biological activity is slightly higher at the L mine site in com-parison with the deciduous site A. Furthermore, the biological activity decreases morequickly with the depth of soil layers at L mine site than at A mine site (Fig. 1).

Table 6. Results of bait lamina tests. Total feeding activity in % for a 10 days period; Ac, At crests and troughs resp. at mine site A; Lc, Lt at mine site L, respectively; P = level of significance, Kruskal-Wallis-H-Test

Site (LTH) Ac At Lc Lt P

October 1997 27.5 16.5 11.0 47.1 0.037*May 1998 22.5 28.5 21.9 32.4 0.161September 1998 20.2 41.9 30.8 47.6 0.029*Mean (%) 23.4 29.0 21.2 42.4

The minicontainer test method was introduced by Eisenbeis (1994) as a refined litterbag test to study the rates of decomposition of a test material under different partici-pation of soil fauna. Preliminary minicontainer tests were established at the four baitlamina test sites from Oct. 20th, 1997 to Jan. 4th, 1999. The decomposition was nearlycompleted in this period in minicontainers with 2 mm mesh size (participation of atleast the whole soil fauna), reaches about 50% in minicontainers with 500 µm meshsize (participation of the meso- and microfauna) and about 40% in minicontainerswith 20 µm mesh size (expected main participation of the microfauna only; Fig. 2).

Soil microarthropods play an important role in decomposing the poplar litter wit-hin the minicontainer bags. They invade the bags in such numbers that up to 130 ani-mals (an average of about 10 animals) were found in one bag with 250 to 125 mg drymass of litter (Table 7). A concentration of microarthropods to such an extent has ne-ver been found in natural litter layers. Therefore we suppose that within the minicon-tainer bags abnormal conditions of decomposition arise. Collembolans invade thebags in by far the highest numbers (79.5% of the total), followed by parasitiforme,trombidiid and oribatid mites as well as small-sized millipedes. Furthermore, the re-sults show that the objective of using different mesh sizes to exclude the macrofauna(by 500 µm mesh size) and the mesofauna (by 20 µm mesh size) has not been fully re-ached as small springtails (predominantly Folsomia candida) settle the 20 µm bags


after some 6 months in very high numbers. All the other groups of microarthropodswere very seldom found in the 20 µm bags and largely prefer the 500 µm bags.

Possibly the settlement of the minicontainer bags by microarthropods can be usedas an indication of decomposition intensity. Up to 24 weeks after exposure the attrac-tiveness of the containers rises significantly for large and medium mesh size types.Only 6 weeks later, very suddenly, a settlement of the 20 µm containers arises mainlythrough collembolan invasion. This leads to a more intensive decomposition in con-tainers of this type and a decrease in microarthropods in containers with medium andlarge mesh size. This is corroborated by the nearly total decomposition of litter in 2mm mesh containers, which is hardly to be seen in the 500 µm mesh containers. Therole of collembolans and their ability the overcome the 20 µm barrier is discussed byDunger et al. (subm.).

Another possibility for estimating the role of soil fauna in decomposing the deadorganic matter is to calculate the potential zootic decomposition level (DLZpot) afterDunger & Fiedler (1997) from the quotient of the metabolic equivalence (ME) and theabove soil net primary production excluding the wood (see methods). The ME of lum-bricids increases continuously at Berzdorf mine sites under deciduous afforestation up

Fig. 1. Mean feeding activity revealed by bait-lamina (0-8cm) from four test sites atthe Berzdorf mining district; test period May 15th to June 4th , 1998. A= deciduous af-forestation; L= primarily pine afforestation; c= crests; t= troughs. Each test site con-sists of three plots of 16 bait-lamina


Fig. 2. Kinetics oflitter mass loss. Mini-containers with diffe-rent mesh sizes, im-planted horizontallyinto the soils (5-8cmcm depth, followingthe Ah horizon) of the46-year-old mine sitesA (deciduous wood)and L (mixed wood) ofthe Berzdorf miningdistrict. Exposed bet-ween Oct. 24th 1997and January 4th, 1999

Table 7. Invasion of soil microarthropods into minicontainers with poplar litter atBerzdorf mine sites A and L during 14 months

numbers of animals with respect to different mesh sizesdate 2 mm 500 µm 20 µm total collembola

01.12.97 67 68 2 137 2712.01.98 74 87 17 178 7823.02.98 74 21 18 113 7506.04.98 99 133 16 149 11518.05.98 155 161 34 350 20229.06.98 226 254 286 766 63610.08.98 185 450 525 1160 105121.09.98 126 182 415 723 61402.11.98 107 197 514 818 74204.01.99 117 157 487 761 642

total 1230 1710 2314 5254 4179ind. container-1 7.7 10.7 14.5 10.5 8.7


to an age of 33 years, but decreases slightly at the 46 year level. At mine site L whichstarted as a coniferous afforestation and changed into a mixed wood within 20 years,the lumbricid ME increases more slowly up to 46 years. At deciduous sites, the MEvalues of microarthropods reach very quickly the highest level (pioneer optimum af-ter 3 years), decrease up to the tenth year and than increase again to a similar level bet-ween 33 and 46 years. At the mixed coniferous site L the microarthropod ME valuesstart at a very high level and rise again up to 33 years but decrease later up to the 46years age (Table 8).

The calculation of DLZpot as shown in Table 8 is incomplete because it is basedon lumbricids and microarthropods only. Furthermore, the saprophagous macrofaunaand the enchytraeids, as well as the microfauna, are expected to make an appreciablecontribution to decomposition. The macrofauna and enchytraeids have not been stu-died with methods adequate enough to be included in the calculation; for the microfa-una there is no experience for doing so. However, as DLZpot values are guidelinedata, i.e. not for drawing up a balance sheet, the values based on lumbricids andmicroarthropods only are sufficient to estimate the development of the faunal potencyin decomposition. The results (Table 8) indicate that the soil fauna needs, under goodconditions (deciduous woodland) some ten years to reach a decomposition efficiencycomparable to natural sites. This process was impeded at the coniferous afforestation(site L) for over 46 years, although it had changed into a mixed woody stand (L46) inthe meantime.

Interrelationships between soil faunal components

Within the process of new formation of habitats after the technical deposition of lig-nite mine spoils there are not only interactions between immigrating animals andabiotic life conditions, but to a great extent interrelationships between species withdifferent abilities to use the actual life situation. In this context, dispersal potencyseems to play a minor role compared to the ecological potency of the species.


Tabl

e 8.

Yea

rly

litte

r pr

oduc

tion,

met

abol

ic e

quiv

alen

t va

lues

of

lum

bric

ids

(ME

Lu)

and

mic

roar

thro

pods

(M

EM

a) a

nd p

oten

tial

zoot

ic d

ecom

posi

tion

leve

l (D

LZ

pot)

of

lum

bric

ids

and

mic

roar

thro

pods

in B

erzd

orf

min

e si

tes

(se

e m

etho

ds; n

umbe

rs b

ehin

d si

tein

itial

s m

ean

the

year

s af

ter

recu

ltiva

tion)

N1

T3

H7

A10

L10

A33

L33

A46

L46

litte

r (g

dw

m-2

a-1 )

3544

232

728

2n.

d.n.

d.n.

d.57

046

8lit

ter

(kJ

m-2

a-1 )

659

8328

6161

5313

n.d.

n.d.

n.d.

1073

988

18M

EL

u0.

00.

3979

.76

401.

3356

.77

901.

9637

7.15

781.

0650

4.66

ME

Ma

5.16

33.2

922

.04

8.91

51.5

825

.98

78.2

622

.84

21.1

5M

EL

u+M

a5.

1633

.68

101.

8041

0.24

108.

3592

7.94

455.

4180

3.90

525.

81M

EM

a : M

EL

u %

100

98.8

21.6

2.2

47.6

2.8

17.2

2.8

4.0

DL

Zpo

t (L

u+M

a)0.

780.

401.

657.

72n.

d.n.

d.n.

d.7.

495.

96

Tabl

e 9.

Rel

atio

nshi

ps b

etw

een

grou

ps o

f m

icro

arth

ropo

ds d

urin

g th

e de

velo

pmen

t of

tw

o m

ine

site

s (A

deci

duou

s w

ood,

Lco

ni-

fero

us d

evel

opin

g to

mix

ed w

ood)

of B

erzd

orf o

pen

cast

min

ing

dist

rict

dur

ing

46 y

ears

. D=

den

sity

(100

0 in

d. m

-2);

% =

dom

inan

cepe

rcen

tage

Yea

r19

6219

6219

8519

8519

9819

98Si

teA

10L

10A

33

L33

A46

L46

Den

sity

/Dom

inan

ceD

%D

%D

%D

%D

%D

%

Col

lem

bola

8.9

36.9

6.5

5.3

27.8

32.9

35.9

15.4

28.2

38.9

27.9

43.6

Prot

ura

1.2

4.9

––

2.5

2.9

2.7

1.2

1.9

2.6

2.4

3.8

Ori

batid

a6.

225

.710

2.0

83.3

26.5

31.3

95.2

40.8

26.8

37.1

20.2

31.6

Tro

mbi

difo

rmes

6.5

26.8

12.2

9.9

18.3

21.6

87.6

37.6

4.9

6.9

2.9

4.6

Para

sitif

orm

es1.

45.

61.

81.

58.

610

.111

.34.

88.

211

.47.

211

.2

Tota

l24

.210

012

2.5

100

84.6

100

233.

210

072

.210

064

.010

0


Table 10. Relationships between genera of lumbricids during the development ofBerzdorf mine sites over 46 years. Age = years after afforestation, mean = mean totalbiomass (g wm m-2), site see „study sites“

Age site mean biomass biomass (%)Aporrectodea Dendrobaena, Lumbricus Octolasion

Dendrodrilus

deciduous af forestat ions2 N 03 T 0.14 1005 N 2.3 1007 H 6.7 44.0 53.0 0 3.010 A 40.6 62.7 10.6 24.6 2.114 A 49.5 27.9 2.3 64.5 5.333 A 104.2 49.2 2.1 47.4 1.334 A 83.2 44.3 1.0 54.7 046 A 83.1 28.6 2.0 58.6 10.8 primari ly coni ferous af forestat ion10 L 4.5 12.7 87.3 0 033 L 38.2 20.4 3.8 74.7 0.934 L 41.3 34.5 2.9 62.0 0.646 L 52.8 19.2 6.1 72.9 1.8

Table 11. Comparison of the developmental stages of lumbricids and microarthropods (measured as respiratory equivalences per m2 = RE, see methods) at Berzdorf minesi-tes 2 to 46 years after the afforestation. Age = years after afforestation, site see „study sites“, n.i. = not investigated

deciduous afforestations primarily coniferous afforestationAge site RE lumbricids RE microarthropods site RE lumbricids RE microarthropods

1 N 0.0 0.56 L n.i. n.i.3 T 0.05 3.92 L n.i. n.i.7 H 9.25 2.80 L n.i. n.i.10 A 47.93 1.12 L 6.78 6.1633 A 107.73 3.36 L 45.05 9.5246 A 95.15 2.96 L 56.71 2.59

With respect to the microarthropods, Collembola, Oribatida and trombidiid mitesare the most important immigrators into young mine sites. Their relative developmentwithin the different mine sites clearly depends on the specific conditions (Table 9).The oribatids dominate at the 10 years old L mine site in the pure Pinus-needle litter,but take second place, behind Collembola, 36 years later in the mixed deciduous andneedle litter at the same site. Even in this case, as well as generally at the A mine site,the proportion of the oribatid density approximates that of the Collembola, which ge-neral dominate at A mine site and at L mine site in 1998. Only the trombidiid mites


have a comparably high share of the density at A mine site in 1962 and 1985 and at Lmine site in 1985. The parasitiforme proportion, esp. Gamasida, is comparable on theolder sites with mull humus (A 33+46, L46), but their absolute abundances were hig-hest at L33 with mixed raw and moder humus. This stage bears the highest microar-thropod density.

The development of the lumbricids at the mine sites A and L is a meaningful ex-ample of the interrelationship of genera within a family (Table 10). Under the condi-tion of nutrition with deciduous litter the detectable settlement of lumbricids started atmine site A in the fourth year with Aporrectodea caliginosa, soon followed by Den-drobaena octaedra. After 10 years the population is dominated by Aporrectodea ca-liginosa (with small amounts of A. rosea), sharing with Lumbricus rubellus (25%)and Dendrobaena octaedra (11%). Only 4 years later the proportions between Lum-bricus (now with L. terrestris as well) and Aporrectodea is reversed, indicating a highadvantage of the nutrition from the surface of the litter layer. Later on the proportionbetween these genera realigned again. The genera Dendrobaena/Dendrodrilus andOctolasion play a minor role during the whole development.

Other conditions prevail at site L with primarily strong raw humus formation. Thefirst coherent populations were built up by Dendrobaena octaedra, which continuesto be dominant (with the participation of Dendrodrilus rubidus) over nearly 30 years,proving the predominant influence of an exohumus layer at this site. After 33 years,and later on, the genus Lumbricus (already with a major proportion of L. terrestris)clearly dominates at site L, too. Up to the last investigation (46 years) the Aporrecto-dea density is relatively small, but clearly greater than that of the genera Dendroba-ena/Dendrodrilus and Octolasion, the latter showing only a minor presence.

The succession of the lumbricid genera and species demonstrated above may begenerally caused by the ecogenesis of the studied mine sites (Dunger & Wanner1999a). Additionally, interrelationships between species and groups of animals playan important role. It may be conceivable that the incorporation of ectohumus into themineral soil layer by anecic earthworms results in deprivation of the epigeic forms oftheir habitat and nutrition.

Even more interesting is the question whether there are interrelationships betweenmembers of the macrofauna and the mesofauna during succession. For quantitativecomparisons, neither population densities nor biomasses are suitable, but the basal re-spiration at 10°C related to the individual weight multiplied by the average biomassper m2 (RE = respiratory equivalence).

The successional comparison in earthworms and microarthropods based on RE va-lues (Table 11) at site A shows that the energetic capacity of the microarthropods (es-pecially Collembola and Oribatida) rises very suddenly during the first three to fiveyears, but is then outstripped and depressed by the development of the earthworm po-pulation and the diminution of the litter layer. The microarthropods need some 20years (more exact intermediate studies are required) to overcome that depression. Du-ring this time a basal change of the communities occurs (Dunger 1991a; Dunger &Wanner 1999b). A retroaction on the development of the earthworm communities cannot be proved from the facts known at present.

The same applies to the primary coniferous site L, but with a retarded influence ofthe earthworms (not earlier than the tenth year) because of the poor palatability of theneedle litter for most lumbricids. Thus, the microarthropods starting on site L with


double the intensity compared to site A, show nearly no reaction against the limitedearthworm activity, because the exohumic situation is altering to a lesser extent andover a longer period than in deciduous humic stands. After 46 years the humus layerseems to get an adequate quality for microarthropods as on site A. That is corrobora-ted as well by the approach of the energetic capacity of the earthworm population atsite L to that at site A in this time.

The presented development of the interrelationships between earthworms andmicroarthropods can be derived as well from the quotients of microarthropod andlumbricid metabolic equivalences (ME) listed in Table 8. At deciduous stands the quo-tient starts with nearly 100%, decreases in the „contact time“ down to 22% and rea-ches a low (2.2%) as soon as after 10 years. At primarily coniferous stands this suc-cession takes more time and comes to a low as late as after 46 years.

Discussion

Mine site features

A detailed knowledge of the geological substrate as well as the type of equipment usedin mining is a prerequisite for comparisons of faunal succession at strip mine dumps.The older Berzdorf spoils had been dumped half a century ago with a primitive tech-nology, thus distinct cover materials supporting reclamation are lacking. Reclama-tion-supporting mining technologies, as are usual at e.g. the Rhineland open-cast mi-ning district (Dworschak 1997; Topp et al., in press), are generally claimed since 1980by the Federal Mining Act as a „bed of arable land“ to the depth of 0.6-1.0 metres oftopsoil (Hildmann & Wünsche 1996). On the other hand, the Berzdorf top soil sub-strates which are mixed materials from very different strata of the overburden, are notas highly contaminated by pyrite and marcasite as those materials used at the CentralGerman or Lower Lusatian mining districts (Hüttl et al. 1996). The severe problemsof sulphur acidification detrimental for plant and animal immigration (Brüning et al.1965; Dunger 1969, 1979; Dunger et al. 1997a, b; Keplin & Hüttl 1999) are not typi-cal for the Berzdorf opencast district.

Another point to bear in mind if comparing mine site development is different hu-man activity. A ‚pure‘ natural succession is described rarely (Jochimsen 1996), inve-stigations on succession on afforestation after amelioration or agricultural utilisationare more usual (Curry & Cotton 1983; Rushton 1986;Wermbter 2000). The Berzdorfmine sites have received only a minor amelioration (mainly liming) and virtually noforestry maintenance since plantation.

An important condition, at least for the first years of renaturation /recultivation, isthe surface quality caused by the dumping technique. The using of stackers or primi-tive railborne techniques with tipper lorries, as used at the Berzdorf dumps, frequentlyresulted in long stripes of crests and troughs. As emphasized also by other authors(Topp et al., in press), Dunger (1968, 1989) found at Berzdorf dumps the earliest sett-lement and development of vegetation and fauna in troughs, virtually based on lowertemperature extremes, a more constant moisture, a finer soil texture and higher con-tents of organic matter. Today, forty years later, the micro-climatic differences bet-ween troughs and crests are less pronounced, because differences in altitude became


equalized by erosion, whereas the higher quality of soil texture and organic matterconcentration in troughs remain to some extent (Dunger et al., in prep.). Thus the eco-logical preference of the soil fauna towards troughs is less pronounced than expected,even though some indication is still given today (Wanner & Dunger, subm.).

Immigration and development of the soil fauna

The up-to-date results on the investigation of the faunal succession in the Berzdorfmining district (Dunger & Wanner 1999a, b;Wanner et al. 1998; Wanner & Dunger1999) lead to the conclusion, that natural immigration by air-, water- , material-trans-port, or active locomotion ensure the settlement of nearly all groups of the soil faunadepending on the dump top soil quality. These studies support earlier observations ofthe „arthropod fallout“ (Crawford & Edwards 1986; Klausnitzer 1993) as an impor-tant source of mine area settlement. Experimental application of (faunated) woodlandtop soil and litter (Glück 1989; Wolf 1985) had, with respect to soil fauna, no clear be-neficial effects. In some cases, the introduction of suitable species of earthworms suc-ceeded in acceleration of soil biological processes, esp. in stabilising aggregates andincreasing soil microbial biomass (Scullion & Malik 2000). However, such effectshave also been shown at dumps without inoculation of earthworms or other compo-nents of the soil fauna, and even are to be demonstrated under the hampering influ-ence of pyrite substances (Keplin et al. 1999; Wermbter 2000). The soil profile inve-stigations (Dunger & Wanner 1999b), as well as the decomposition study (Figs. 1, 2),confirm an equivalent development of the soil biological processes at the Berzdorfmine sites.

This conclusion is corroborated by the quantity and species composition of the soilfauna. The earthworm communities established on the Berzdorf mine sites are com-parable with those of the forest reclamation areas of the Rhineland mining district(Topp et al. 1992), as well as with different tree plantations (Quercus, Tilia: Topp etal., in press; Dworschak 1997, with additional Lumbricus castaneus), under agricul-tural reclamation (without epigeic earthworms, Westernacher-Dotzler & Dumbeck1992), or even with colliery spoil heaps at Durham (additionally with Lumbricus fest-ivus, Standen et al. 1982). As this can be expanded to communities from North Ger-man fallows (Bessel & Schrader 1998), as long as the area of distribution of peregrinespecies only (Brauckmann et al. 1995) is not exceeded, it seems that it concerns a„community of disturbed areas“. The long-term observations on Berzdorf mine sitesconfirm a varying presence of species such as Allolobophora chlorotica or Aporrec-todea longa, a process that corroborates the conditions postulated by den Boer (1981)for heterogeneous and variable life situations. Only the unstable participation of Oc-tolasion lacteum is not understandable by these reflections. In this context, it shouldbe examined to which extend the soil genetical circumstances are realised as deman-ded by Höser (1994) for settlements of this species. The well known influence oflong-term changing of climatic factors on the earthworm species spectrum (Scheu1992) was not recorded in our studies, because the samplings during a 2-3 years pe-riod were not dense enough. Similar assessments, as demonstrated for earthworms inthe present study, can be easily applied to other groups of the soil fauna (Dunger &Wanner 1999b).

In spite of the importance of soil Protozoa and Nematoda with respect to generalecology and bioindication (e.g. compiled in Darbyshire 1994; Freckman 1982), most


studies on mine site recultivation and succession have not considered the soil micro-fauna at all. As shown in our study, microfaunal (e.g. testate amoebae and nematodes)diversity and community development present valuable information about the statusof soils in ecogenesis. Testate amoebae as well as nematodes occur early (within a fewweeks) on freshly deposited mine dumps (Wanner et al. 1998), developing into site-specific populations within a short time span. Compared to plant succession of spoilsoils, the changes in the nematode communities occur considerably faster. ThereforeBongers & Bongers (1998) argue that „each characteristic successional stage offersthe nematode fauna enough time to develop towards the climax community that is ty-pical for that vegetation type“. In conclusion, expanding our knowledge on microfau-nal primary succession (e.g. on coastal dunes: Goralczyk & Verhoeven 1999), it be-comes obvious that the role of the microfauna has definitely to be considered in furt-her studies on ecosystem functioning and primary succession.

Action of the soil fauna in ecogenesis of mine sites

Due to determination problems of soil organic matter (SOM) caused by a varying lig-nite content of the spoil substratum, continuous studies on SOM are not available.Comparing the soil quality of the first years of succession (Dunger 1968) and after 45years (Hauser & Kowarsch 1998; Kobel-Lamparski & Lamparski 1999), it can clearlybe concluded that the profiles of the studied mine sites are structured by earthwormactivity. In contrast to the findings of Topp et al. (in press) from Rhineland spoils, thepH values remained at the same level or even decreased slightly (Table 1). The influ-ence of the growing woodland can be derived from the litter production (Table 8). Asstated by other investigations from mined areas (Felinks et al. 1999; Wiegleb et al.2000), the developing of the vegetation is the main factor for changes in species com-position.

Table 10 shows the varying contribution of earthworm species at the Berzdorfmine sites. The participation of the epigeic Dendrobaena octaedra mainly during thefirst years at mine site L can not only be seen as an indication of a strong litter layerupon the mineral soil but also as a distinct qualitative influence on decomposition.Scheu & Parkinson (1994) found that this species increases the leaching of nutrientsand reduces the microbial biomass at least in decomposing N-rich aspen litter. As forthe dominating anecic Lumbricus terrestris, recent studies (Daniel 1991) corroboratethe old knowledge (Dunger 1964) of a high leaf-litter consumption.

The development of the earthworm community at the studied mine sites may indi-cate the biological soil condition. Using strategy-types proposed by Graefe (1997),the predominating combination between anecic and mineral forms with only a fewepigeic ones is typical for fresh to wet loamy soils. Taking this into consideration,spoil soils (with the minor influence of pyrite or marcasite) are little different from na-tural ones, at least for the meso- and macrofauna.

The contribution of soil animal groups other than earthworms in the developmentof soil structures, decomposition and nutrient cycling is also obvious from the resultsbut less discernible. At the end of the pioneer optimum (7th to 12th year at mine siteA; Dunger 1989) the increasing activity of anecic earthworms (Lumbricus terrestris)significantly affects the ectohumic litter layer and thus destroys the habitat of most ofthe microarthropods (esp. oribatid mites and springtails). In the primary coniferous


site L the process went on for about twenty years. Additionally, a pronounced decreasein density of these groups occurred (Table 9), thus resulting in an obviously negativeeffect on the earthworm populations. On the other hand, from the burrowing activityand the humic walls in the drilosphere, a positive influence might have been expected(Tiunov & Scheu 1999). Direct interrelationships between earthworms and microar-thropods, as shown by Scheu et al. (1999) in experimental systems, indicate differen-ces in the reaction of edaphic and epedaphic Collembola. Correspondingly, the suc-cession process on Berzdorf mine sites proceeds independently for edaphic and eped-aphic communities of Collembola (Dunger 1991a). The details of the transfer of ex-perimental results into field conditions, especially concerning the incorporation of theterrestrial detritivore system into the complex trophic interactions (Gange & Brown1997), are still poorly understood, insufficient for more far-reaching presumptions.The same applies to presumable interactions (Alphei et al. 1996) between Protozoaand Nematoda with the soil meso- and macrofauna. More details will be presented inDunger et al. (in prep.).

The decomposition tests (bait lamina, minicontainers) were expected to show ahigher biological activity at the deciduous mine site A. This could not be proved. Sur-prisingly, the minicontainers not only exhibited an extreme „litterbag effect“ (Cros-sley & Hoglund 1962), but also the highest collembolan density in the containers with20 µm mesh size which were expected to be free from mesofauna. It is important toknow that these high densities occurred as late as 7 months after starting. After thistime Folsomia candida, being nearly absent at samples from soil cores, invades thecontainers filled with poplar litter up to 600 specimens per g dry mass. It is supposedthat newly hatched instars had been attracted by the decomposing litter and were ableto penetrate the gauze, but after increasing in body size could not escape again (Dun-ger, Schulz & Zimdars, subm.). Keplin (1999) found at reclaimed pine mine sites withthe same technique, but with Pinus sylvestris root litter and a shorter observation time,55 specimens per 1 g litter at maximum. However, assessing decomposition by thismethod may not be suitable to reveal values typical for the investigated stand (Törne1997).

The meaning of soil fauna for a „good restoration“

Finally, it may be asked for the meaning of the soil fauna in the discussion about„good ecological restoration“ (Higgs 1997), nowadays considered in a broad context(Hüttl et al. 1996, 1999; Pflug 1998; Broll et al. 2000; Wiegleb et al. 2000; Topp et al.,in press). The goal is a self-sustaining rehabilitation (Langkamp et al. 1979), finallyreaching the „original ecosystem“, i.e. a full restoration approaching the „preturbationpoint“. This is seldom achieved and the more frequent alternative approach is the re-placement by another system (Majer 1989). The importance of the soil fauna in theprimary succession process to improve the soil structure and enhance decompositionand nutrient cycling is briefly described in the present case study „Berzdorf mines“.The questions remain whether a typical, „preturbation“ composition of the soil faunahas to be demanded or even claimed, as it is in case of the vegetation cover or birdcommunity, to fulfill the claims for a „good ecological restoration“. With the progressin soil zoology, this goal is not an illusion, but rather a possibility to proof the self-su-staining condition of the ecosystem. We know not only for the earthworms (Graefe1993) but more precisely for microarthropods (Dunger 1991a; Beck et al. 1997) that


„decomposer communities“ with regional differentiations may have characteristicspecies compositions depending on soil properties such as texture, moisture etc.(Römbke & Dreher 1999). Thus, we can conclude that the soil faunal development atthe studied mine sites of Berzdorf fulfills the demands for a good ecological restora-tion at least for the deciduous site A, despite a very bad and irregular mineral compo-sition in the cover layer of the dump. In contrast, our investigations on a 35-year-oldpyrite-rich lignite spoil at Domsdorf (Lower Lusatia), afforested with Quercus rubraand Tilia cordata, resulted in a very poor soil fauna, partly caused by planting exoticoaks and exhibiting a „bad ecological restoration“ (Dunger 1997b).

Acknowledgements

We thank Dr. Beate Keplin for supporting our decomposition study. Prof. Dr. W. Xylander en-couraged us with beneficial discussions and institutional help. The technical assistance of Ker-stin Franke, Marlis Römer, and Heiderose Stöhr is gratefully acknowledged. This study was fi-nancially supported by the BMBF (#0339668; the responsibility for the content of this publi-cation is taken by the authors).

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