Management

Forest Ecology and Management 77 (1995) 35-51

The effects of clear-cutting and liming on the soil macrofauna

of a beech forest
Anne Theenhaus * , Matthias Schaefer
II. Zoologisches Instinct, Abteilung Ckologie, Universitiit GBttingen, Berliner Strasse 28, D-37073 Giittingen, Germany

Accepted 1 May 1995

Abstract

The effect of clear-cutting and liming on abundance, dominance structure and trophic composition of the soil macrofauna
(Lumbricidae, Diptera, Coleoptera and Chilopoda) was studied in a beech wood forest on an acidic brown-earth. Sampling
methods were ground photo-ecledors and funnel extraction. The various taxa and nutritional groups of Diptera responded
differently to the environmental changes caused by clear-cutting and liming. The same was true for the species and taxa of
Coleoptera. Lumbricidae and Chilopoda were favoured by the liming treatment. The factors determining the change in
population densities are discussed. Abiotic factors, especially soil moisture, and biotic factors, such as dead plant biomass
and predation, are considered to be of prime importance. A change of moder soil into mull soil after liming is observed.

Keywords: Clear-cutting; Liming; Beech forest; Macrofauna

1. Introduction macrofauna from the initial state of a succession

Clear-cutting as a restocking method is common
practice in today’s forestry management. In addition,
the liming of forests is a common strategy for miti-
gating the problem of acid rain. However, the envi-
ronmental effects of these treatment strategies and
the interaction between them are not well under-
stood. It is quite reasonable to assume that abrupt
changes in an ecosystem, such as clear-cutting and
liming, would be mirrored in the composition of the
animal community. The objective of this study is to
determine the changes in population density, domi-
nance structure and trophic composition of the soil
* Corresponding author.
after clear-cutting, with and without liming.
According to Odum (19711, in the early stage of
ecological succession community respiration is low
compared to mature ecosystems. Schaefer (1991a)
points out that decomposition is less than net produc-
tion during the development of a forest. The number
of decomposers is consequently lower than in mature
stands. In this study it is, therefore, hypothesized,
that the decomposer community declines after clear-
cutting.
Furthermore, we assume that the saprophagous
macrofauna is more abundant on the limed plots and
causes a change of moder into mull profile. Schauer-
mann (1986) states that, in general, saprophages are
more abundant in mull soils than in moder soils.
According to Schaefer (1991b), decomposition rates
are generally higher in mull than in moder or mor

037%1127/95/$09.50 0 1995 Elsevier Science B.V. All rights reserved

SSDI0378-1127(95)03580-X
36 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51

forest soils, contributing to a lower standing crop of

soil organic matter in the first soil humus form. In
mull soils, the higher decay rates are the result of
litter comminution by the macrofauna, mixing of
organic material with the mineral soil and the main-
tenance of well-drained aerobic conditions in the
soil. Moder soil, on the other hand, has low pH
values in all strata, which inhibit the macrofauna and
prevent its feeding and burrowing activities (Swift et
al., 1979).

2. Materials and methods

2.1. Site

The Soiling is a plateau of triassic red sandstone

(Buntsandstein) about 50 km north-west of Gottin- T
gen (Lower Saxony). It is overlain with a layer of 30m clear-cut area
loess loam that is more than 50 cm thick. The soil is I 0 .I
composed of an acidic brown-earth with a pH value
(KC0 lower than 4.2. The humus form is a typical
moder. Since the process of litter decomposition is
I
70m limed area

rather slow, the litter-humus layer is up to 5 cm

thick. The experiment was performed in the upper
Solling (‘Hochsolling’), 500 m above sea level. At
this altitude, the climate is suboceanic-montane (El-
lenberg et al., 1986). Average annual rainfall is 1045
mm, average annual temperature 6.5”C.
The site under investigation, previously desig-
nated as B2 plot, is a beech forest of the Luzulo-
Fagetum typicum type. Most of the trees, predomi-
nantly Fagus syluatica L., are about 150 years old.
The ground flora consists of Luzula luzuloides
(Lam.), Oxalis acetosella L., Deschampsia flexuosa
L. and seedlings of Fagus sylvatica L. The ground
flora is extremely sparse, and a shrub layer is not
present. A detailed description of the study site is
given in Ellenberg (1971).
2.2. Experimental design
In October 1989, four clear-cuts, measuring 30 m
in diameter, were created at least 50 m apart. A total
of 64 beech trees were cut and removed from the
experimental site. Two of the clear-cuts, including a
surrounding forest area 20 m wide, were limed with
fine-grained magnesium carbonate in a concentration
I 0 /’
Ir zoological plots
(1Om x 5m)
Fig. 1. Experimental site (B2 area) in the beech forest of the upper
Solling (Lower Saxony).
of3tha-‘. Eight plots, each measuring 10 m X 5 m,
were reserved for zoological research. Four of them
were located within the clear-cuts, two of which
were limed. The other two plots in the clear-cut areas
remained unlimed. Each of the other four plots were
located south of a clear-cut area within the beech
forest. Two of the plots were located within the
limed areas, the other two in the unlimed areas (Fig.
I).
2.3. General properties of the tqwrimental plots
2.3.1. Vegetation and litter fall
The limed plots had a dense herb layer. In June
1992 it covered 70-100% of the forest floor. The
dominating species was Epilobium angustifolium
CL.), followed by Luzula Zuzuloides(I&RI.) and Ox-
 A. Theenhaus, M. Schaefer/ Forest Ecology and Management 77 (1995) 35-51 37

Table 1
pH and element concentrations of the organic horizons O,, Or and 0, in the beech forest stand, the unlimed and the limed clear-cut plots
Horizon pH (CaCl,) P K Ca Mg
k, iz8 (mg g-l) (mg g-l) (mg g-l) (mg g-l)
Unlimed clear-cut
0, 3.42 b 43.3 1.72 0.94 1.83 3.92 b 1.11
Of 2.81 b 34.1 ab 1.93 ab 0.95 1.37 b 2.31 b 0.97 b
Oh 2.72 b 31.3 1.71 1.15 a 3.65 1.07 b 1.86 ab

Limed clear-cut
0, 4.79 a 41.5 1.58 0.98 2.24 7.08 a 2.18
0‘ 4.11 a 34.2 b 1.60 b 0.84 2.91 a 12.16 a 4.89 a
Oh 3.38 a 30.6 1.62 0.69 b 3.16 3.39 a 3.89 a

Beech forest stand

0, 3.83 ah 43.7 1.77 1.05 2.01 4.86 ab 0.75
Of 2.88 ab 43.7 a 1.99 a 0.85 1.40 b 2.81 ab 0.58 b
Oh 2.69 b 32.1 1.70 0.98 ab 4.12 1.15 b 1.20 b

Figures sharing the same letter are not significantly different (Wilcoxon-test at P < 0.05) (Bauhus, 1994).

alis acetoselh L. By contrast, the unlimed plots were 2.3.3. pH and element fluxes
sparsely covered with herbs (2-20%). pH, element concentrations and element exchange
Leaf litterfall into the clear-cuts from the sur- capacity are presented in Tables 1 and 2.
rounding beech trees amounted to 75% of the litter- In the beech forest stand negligible amounts of
fall in the beech forest (2.9 t ha-‘). Total litter input nitrate were measured in the seepage water at 80 cm
in limed clear-cuts increased, because on these plots depth (Bauhus and Bartsch, 1995). Here sulphate and
an additional 2.8 t ha-’ of organic dry matter was aluminium were the dominant ions in the seepage
supplied by the above ground herbaceous vegetation water. In the clear-cut plots disruption of plant up-
(Bauhus and Bartsch, 1995). take and high infiltration rates in the unlimed clear-

2.3.2. Precipitation, soil moisture and temperature

Due to the lack of tree cover, in each of the Table 2
clear-cuts, more precipitation reached the ground. In pH and cation exchange capacity of the surface mineral soil in the
beech forest stand, the unlimed and the limed clear-cut plots
1991 and 1992, Bauhus and Bartsch (1995) reported
7% more precipitation reaching the ground in the Depth pH C N K Ca Mg AL
(cm) (CaCl,) (%I (o/o) (pm01 IE g-l)
clear-cuts than in the beech forest.
Soil water tension differed enormously between Unlimed clear-cut
o-1 2.76 ab 11.3 0.6 3.4 6.8 b 2.9 77.2
the clear-cut plots and the beech forest plots. In the
l-2 2.83 ab 6.1 0.3 2.9 ab 4.7 b 2.1 82.9
clear-cut plots, the water tension at a depth of 15 cm 2-5 2.94 ab 4.4 0.2 2.2 b 2.8 b 1.3 88.1
was close to saturation point throughout the year. By S-10 3.17 ab 3.4 0.2 1.8 b 2.2 a 1.1 98.5
contrast, in the beech forest the soil dries during late
Limed clear-cut
summer (830 hPa), because of water removal by the o-1 3.19 = 10.9 0.6 2.9 36.1 a 15.3 52.7
beech trees (Bauhus and Bartsch, 1995). l-2 3.03 = 7.1 0.4 2.0 a 14.6 a 6.5 72.0
Average daily soil temperatures in the clear-cuts 2-5 3.07 a 4.3 0.2 1.5 a 8.3 a 3.4 82.7
were not different from those in the beech forest S-10 3.30 a 3.0 0.2 1.2 a 3.2 a 2.8 87.7
Beech forest stand
stand on average. However, maximum daily temper-
o-1 2.71 b 12.0 0.6 4.3 4.0 b 2.5 73.4
atures were on some occasions warmer in the stand l-2 2.78 b 6.5 0.3 2.9 b 2.8 b 1.8 83.3
than in the clear-cuts (Bauhus and Bartsch, 1995). 2-5 2.89 b 4.3 0.2 2.0 ab 1.8 b 1.2 93.7
This was most obvious during the summer when S-10 3.15 b 3.1 0.2 1.4 a 1.2 b 1.0 96.9
maximum temperatures at 5 cm depth were some- Figures sharing the same letter are not significantly different
times 4°C higher than in the beech forest stand. (Wilcoxon-test at P < 0.05) (Bauhus, 1994).
38 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (199.5) 35-51

cut plots caused substantial nitrate losses: 80 kg Table 3

NO;-N ha-’ aa1 were lost from the unlimed clear- Population density (ind. m ’ ) of Lumbricidae on the limed (Ii)
and unlimed (unli) clear-cut and the limed (ti) and unlimed funli)
cut plots and 30 kg NO,-N ha- ’ a-r from the limed beech forest plots in the Solling mountains (1992/1993)
clear-cut plots (1992) (Bauhus, 1994). This process ____-. _-
Clear-cut Forest
caused soil acidification and aluminium release.
li unli Ii unli
Therefore, nitrate and aluminium were the dominant -- -~.-
ions in the seepage water of clear-cuts. Dendrobaena octaedra 6av.i 19 3 16 i
Dendrodrilus rubidus (Sav.) 20 1 20 0
In the limed clear-cut plots (1991), less nitrogen
Dendrobaena / Dendrodrilus, juvenile 250 7 264 h
was mineralized, compared to the unlimed clear-cut Lumbricus eiseni (Lev.) 9 0 ! 0
plots (Bauhus and Barthel, 1994). Lumbricus rubellus Hoffm. 2 (1 1 !i
Lumbricus, juvenile 8 + 3 ii

2.4. Sampling of the fauna Funnel extraction method (mean of four sampling dates 1992). ;
between 0 and 1 ind. m-l found.

On 11 March 1992, a total of 24 ground photo-

eclectors (three on each zoological plot) (Funke,
1971), each covering 0.3 m2 were randomly set up.
Sampling was done every 2 weeks. The last day of
sampling was 4 November 1992.
Nine soil cores (diam. l/28 m2, 10 cm depth) per
plot, including the litter layer, were taken four times
during that year: in spring (27 May 1992), in sum-
mer (5 August 1992), in autumn (11 November
1992) and in winter (3 February 1993). The samples
were extracted using an apparatus designed by
Kempson et al. (1963), modified by Schauermann
(1982).
2.5. Statistical treatment
The influence of liming and clear-cutting on the
soil macrofauna and the interaction between these
factors were tested by two-way analysis of variance
(ANOVA). A block design was set up, each adjacent
forest plot and clear-cut plot representing one block.
Factors were liming and clear-cutting. Variance ho-
mogeneity was tested using the Bartlett test.
For many groups of soil fauna, caught by the
funnel extraction method, the analysis of variances
was not applied, because there was no variance
homogeneity. In these cases only tendencies are
described.
3. Results and discussion
During the experiment, a total of 1600 specimens
of Lumbricidae, 18 800 of Diptera, 2800 of
Coleoptera, 500 of Lithobiomorpha, 70 of tipulid
larvae and 2400 larvae of Athous subfuscus were
identified. Other macrofauna groups were repre-
sented only with low numbers and were therefore not
included in the study: 12 Gastropoda, 14 Diplopoda,
about 100 Hymenoptera and 15 Lepidoptera.
3.1. Lumbricidae
Liming led to an increase of the abundance of all
lumbricid species (Fig. &a) and Table 3). On the
unlimed plots the average density of Lumbricidae
was 10 individuals (ind.) m-‘, whereas a very high
number, an average density of 307 ind. m-‘, was
found on the four limed plots. A positive correlation
between the density of Lumbricidae and the lime
content of the soil has been found by many authors
(e.g. Piearce, 1972; Hnhta, 1979; Schauermann,
1986; Funke, 1991). Presumably the comparatively
high pH value (Table 1 and 2) of the soil on the
limed plots caused the increase in Lumbricidae. An-
other favouring factor on the limed plots presumably
was the high herbaceous biomass, especially decay-
ing material of Epilobium angustifolium.
Clear-cutting had no obvious influence on the
density of Lumbricidae on both unlimed and limed
plots. Huhta (1976) found an increase in Lumbrici-
dae during the first few years after clear-cutting. The
author attributed this finding to the higher amount of
organic matter remaining in the soil in the form of
felling residues. In the present study, however, trees
were removed from the clear-cuts and, furthermore,
the concentration and cation exchange capacity of
 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51 39

nutrients, such as N and P, did not significantly emergence abundance value of 4114 Diptera rnm2.
increase in the clear-cuts (Tables 1 and 2). Of these 3.3% were Brachycera. Sciaridae were pre-
The occurrence of Lumbric~ eiseni (Lev.) is dominant, comprising 77.9% (3205 ind. me21 of the
worth mentioning. This species is very rare in Ger- total catch, followed by Sciophilidae with a percent-
many. Lumbricus eiseni was found only on the age of 9.5% of the total catch (390 ind. ms2>.
limed plots (both forest plots and clear-cut plots). Cecidomyiidae (including Lestremiinae) comprised
We cannot imagine how this population has colo- 7.1% (292 ind. mp2) of the total catch of Diptera.
nized these plots. Hovemeyer (1991) pointed out that, in acidic beech
woods, Sciaridae are usually more abundant than
3.2. Diptera Cecidomyiinae. The high emergence abundance of
Species belonging to 33 families of Diptera were Chironimidae on the unlimed clear-cut plots, found
found. Of these, 14 belonged to Nematocera and 19 in this study, could possibly be attributed to the fact
to Brachycera. that the emergence abundance of Diptera is subject
The average emergence abundance of Diptera was to fluctuations from year to year (e.g. Altmtiller,
2600 ind. m- 2. The lowest density (1027 ind. mm21 1979).
was recorded on an unlimed clear-cut plot. The Despite the fact that the dominance structure of
highest density (5051 ind. rnp2) was recorded on a the dipterous community varied greatly between the
limed clear-cut plot. Of the total catch 7% were experimental plots, no pattern became evident. Chi-
Brachycera. ronomidae were an exception, since the dominance
The most abundant taxa were Cecidomyiinae position of Chironomidae on the two unlimed beech
(28.7-64.3% of the total catch), Sciaridae (20.1- forest plots was relatively high. However, it is not an
33.8%) and Chironomidae (1.9-17.5%) (Table 4). increase of Chironomidae, but rather the reduced
On all but one plot, Cecidomyiinae were predomi- emergence abundance values of Cecidomyiinae, on
nant, followed by Sciaridae. On the unlimed clear-cut these plots that could be the explanation for this
plots, Chironomidae were relatively dominant com- finding.
prising 17.5% of the total catch. In Table 5, only those Diptera taxa are listed
On an area of beech forest adjacent to the site where the statistical test revealed significant differ-
examined in this study, Altmiiller (1976) found an ences in the emergence abundance between the treat-

Table 4
Cumulative emergence abundance of Diptera (ind. m -‘) and percentage of taxa of the total catch of the respective plot (%, mean values of
two plots) on the limed and unlimed clear-cut and beech forest plots in the Solling mountains (1992/1993) (catches of ground
uhoto-eclectors)
1

Cumulative emergence abundance (ind. m-’ ) Percentage of taxa

Clear-cuts Forest Clear-cuts Forest
li unli li unli li unli li unli
Cecidomyiinae 2373 414 1477 1477 64.3 28.7 51.4 60.8
Sciaridae 742 467 970 589 20.1 32.4 33.8 24.2
Chironomidae 69 253 102 49 1.9 17:s 3.5 2.0
Sciophilidae 73 62 77 85 2.0 4.3 2.7 3.5
Phoridae 118 57 58 47 3.2 3.9 2.0 1.9
Ceratopogonidae 104 64 11 10 2.8 4.4 + +
Lestremiinae 34 14 38 39 + 1.0 1.3 1.6
Empididae 9 32 16 44 + 2.2 + 1.8
Fanniidae 31 14 31 14 + 1.0 1.1 +
Hybotidae 22 19 14 23 + 1.3 + +
Dolichopodidae 29 24 14 6 + 1.7 + +
Scatopsidae 13 0 34 5 + + 1.2 +

Only taxa which comprised at least 1% of the total catch on at least one plot are presented. +, numbers < 1%.
40 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51

Table 5 0.01) (Fig. 2(a)). The same result was obtained for
Taxa of Diptera whose emergence abundance was significantly Cecidomyiinae, which made up 50% of Diptera (P
influenced by the factors of clear-cutting and/or liming (ANOVA)
< 0.01). During the year under observation, two
Taxon Factor Effect Significance emergence peaks of Cecidomyiinae were recorded:
Cecidomyiinae the first in April, the second between the beginning
Total Liming X clear-cutting + r f
- L* * of June and the end of August (Fig. 2(b)). During the
First peak Clear-cutting
Second peak Liming Xclear-cutting + - * * second peak, the highest number of Cecidomyiinae
.I* were detected on the limed clear-cut plots (P <
Ceratopogonidae Clear-cutting +
,.
Dolichopodidae Clear-cutting + 0.001). The April catches were significantly higher
Empididae Liming -
I * * on the beech forest plots, in comparison to the
Fanniidae Liming +
t .I clear-cut plots (P < 0.0011. The reduced emergence
Limoniidae Clear-cutting +
Limoniidae Liming +
*.* characteristic for the first peak on the clear-cut plots
Lonchopteridae Liming Xclear-cutting + * * * is difficult to interpret. The larvae of the Cecidomyi-
. * *
Scatopsidae Liming + inae present are assumed to be gall-making. In au-
4*
Sciaridae Liming + tumn the larvae, inside the leaves, fall to the ground
X , interaction of the factors; + / - , positive/negative effect on and enter the soil for aestivation and/or hibernation.
the emergence abundance. * * *P < 0.001, * * P < 0.01, ’ P < Since all experimental plots had roughly the same
0.05. amount of litter, the decline in the emergence abun-
dance on the clear-cut plots can not be attributed to a
reduced litter biomass. One possible explanation for
ments. The emergence abundance value of total the decline in Cecidomyiinae on the clear-cut plots
Diptera was significantly higher on the two limed could be, that, for some reason, the adult Cecidomyi-
clear-cut plots, compared to the other plots (P < inae avoid the edges of the clear-cuts for oviposition.

Ind /m;
4oc0,
b) Cecidomyiinae
3000

2000

1000

0L -

Ei Nemalocera 0 Brachycera

d) microhumiphagous
Diptera

Fig. 2. Emergence abundances and/or activity density of the soil macrofauna (ind. m-‘) on limed and unlimed clear-cut plots, and on
limed and uniimed beech forest plots in the Solling mountains, sampled with ground photo-eclectors. Capture period 11 March to 4
November 1992. cl noli, clear-cut, not limed, cl li, clear-cut, limed; fo noli, beech forest, not limed; fo li, beech forest, l&ted.
 A. Theenhaus, M. Schaefer/Forest
3.2.1. Trophic groups of Diptera
In order to point out functional aspects, catches of
Diptera were subdivided into trophic groups, accord-
ing to the feeding habitats of their larvae. The arti-
cles by Brauns (19541, Raw (19671, Hennig (19681,
Healey and Russel-Smith (19711, Jacobs and Renner
(1988) and Hovemeyer (1985, 1992) were helpful
literature for this categorization.
The statistical analysis for mycetophagous feeders
(Platypezidae, Mycetophilidae, Sciophilidae and
Lestremiinae) revealed no significant differences be-
tween the treatments. Okland (1994) compared the
species richness of Mycetophilidae between clear-
cuts, managed forest sites, which were clear-cut
70-120 years ago, and semi-natural forests (only
selectively cut). The author found no significant
difference between clear-cuts and managed forest
sites; however, the average number of species was
significantly higher in semi-natural forests. The au-
thor concluded that the restoration of a diverse myce-
tophilid fauna after clear-cutting requires more time
than 70-120 years. This view fits to the findings of
the present study, since the beech forest under survey
is about 150 years old and the mycetophilid fauna
might still be in a developing state.
The necrophagous feeders (Phoridae, Sarcophagi-
dae, Sphaeroceridae) were not influenced signifi-
cantly by the treatments clear-cutting and liming.
The trophic groups showing differences were
macrohumiphages, microhumiphages, ‘surface scrap-
ers’ and zoophages.
3.2.1.1. Macrohumiphages. Those taxa which feed
on litter (microflora and amorphic humus included)
are assigned to the macrohumiphagous feeders. These
are Limoniidae, Trichoceridae, Tipulidae, Scatopsi-
dae, Sciaridae, Bibionidae and Psychodidae.
The emergence abundance of macrohumiphagous
feeders significantly increased after liming (P <
0.001) (Fig. 2(c)). This was mainly due to an in-
crease in population density of Sciaridae (P < 0.011,
Scatopsidae (P < 0.01) and Limoniidae (P < 0.001)
(Table 5). Besides, limoniid numbers were signifi-
cantly higher on the clear-cut plots relative to the
beech forest plots (P < 0.001) (Table 5).
With one exception, no Tipulidae hatched on the
unlimed plots. The emergence abundance value was
2 ind. mm2 on average on the limed clear-cut plots.
Ecology and Management 77 (1995) 35-51 41
Similar patterns were observed for the tipulid larvae
collected by funnel extraction. The highest number
of tipulid larvae were found on the limed plots,
ranging from 8 to 13 ind. m-‘. On the four unlimed
plots 2-7 ind. me2 were found.
The increase in macrohumiphagous feeders on the
limed plots could be attributed to the relatively high
biomass of herbs (especially Epilobium angusti-
folium), which served as food resource for the
macrohumiphagous feeders. In addition, the rela-
tively high pH value on the limed plots could have
favoured the macrohumiphagous feeders, of which
Tipulidae are known to avoid acidic habitats. Irmler
and Heydemann (1989) found a positive correlation
between the pH value of the soil and the average
population density of tipulid larvae. According to
Funke (19911, however, the emergence abundance of
Sciaridae did not change significantly after the appli-
cation of lime and fertilizer to a spruce forest in
Germany.
According to many authors, macrohumiphagous
larvae thrive on moist substrates. Cramer (1968)
observed this preference in the adults and the larvae
of Tipulidae and Limoniidae. Funke (1991) found
that Limoniidae are most abundant in moist forests.
Hovemeyer (1985) reported a decline in emergence
abundance of Limoniidae in years following dry
summers. The development of sciarid larvae is
favoured by a high moisture content of the litter
layer and the upper soil layers during summer dry-
ness (Hovemeyer, 1985, 1991). In this study, how-
ever, no positive correlation between the moisture
content of the habitat and the abundance of macrohu-
miphagous feeders could be found. Instead, this study
revealed an increase in the macrohumiphagous
dipteran numbers on the limed plots. On these plots,
the moisture content did not show an increase at a
depth of 4 cm (the majority of macrohumiphagous
dipteran larvae inhabit a depth of about 4 cm;
Hovemeyer, 1984).
3.2.1.2. Microhumiphages. Microhumiphages feed on
microfungi, algae, Testacea and amorphic humus.
Ceratopogonidae and Chironomidae are assigned to
the microhumiphagous feeders.
The emergence abundances of all microhu-
miphagous feeders (Ceratopogonidae and Chirono-
midae) increased significantly after clear-cutting (P
42 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51

< 0.05) (Fig. 2(d)). Statistical proof for this pattern
could be given for Ceratopogonidae (P < 0.001;
Table 5), but not for Chironomidae. Liming caused
an increase in the emergence abundance of all micro-
humiphagous feeders but a decrease on the beech
forest plots. However, the interaction of clear-cutting
and liming was not significant (P = 0.098).
The significant increase in microhumiphagous
feeders due to clear-cutting could be attributed to the
moisture content of the habitat, as Ceratopogonidae
and Chironomidae prefer moist habitats. The positive
influence of moisture on Ceratopogonidae was
pointed out by Feldmann (1992). Hovemeyer (1985)
reported a positive linear correlation between the
density of ceratopogonid larvae and the relative wa-
ter content of the organic soil layer. Healey and
Russel-Smith (1971) and Heynen (1988) described
the same preference for Chironomidae. It is not
surprising, therefore, that Ceratopogonidae and Chi-
ronomidae preferred the clear-cut plots, because here
the upper strata of the soil, which are the habitat of
populations belonging to these families (Hovemeyer,
1984), were relatively moist compared to the beech
forest plots. During the summer, the water tension in
the beech forest at a depth of 5 cm reached a value
of 600 hPa (J. Bauhus, personal communication,
1993).
3.2.1.3. ‘Surface scrapers’. The ‘surface scraping’
Diptera feed on microorganisms, pollen and humus
material on the leaf surface. The larvae scrape this
material off the leaf surfaces with their mandibles.
Lonchopteridae, Lauxaniidae and Fanniidae belong
to this feeding group.
The emergence abundance values of the ‘surface
scrapers’ increased after liming (P < 0.001) (Fig.
3(a)). The same is true for Fanniidae (P < O.OOl>,
which comprised 65% of the ‘surface scrapers’. Lon-
chopteridae were significantly more abundant on the
limed clear-cut plots compared to all the other plots
(Tukey’s least significant difference of P < 0.001).
Individuals of this family almost exclusively emerged

Ind lm2 ind lmr

ICC!
b) zoophagous Ditm

q Faniwdae 0 Lauxanlldae 0 Lonchoptendae q Empldidae GDollchopod~dae q Hybotldae mother

Ind lm2 lnd /m2

300, 100
r7lI I

I
c) totalCoklptera
d) Curculionidae
200 El

102

q Phyllobius argentatus h3Str melanogrammum

0 eclector method q funnel method q Poh/drusus undatus nother

Fig. 3. Emergence abundances and/or activity density of the soil macrofauna (ind. m -*) on limed and unlimed clear-cut piots, and on
limed and unlimed beech forest plots in the Solling mountains, sampled with ground phot+eclectors. Capture pried 11 Mz&h to 4
November 1992. cl noli, clear-cut, not limed; cl li, clear-cut, limed; fo noli, beech forest, not limed; fo li, beech forest, limed.
 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51 43

from these plots. Emergence abundance values of the ‘surface scrapers’ responded differently to the envi-
Lauxaniidae, however, were comparable for all ex- ronmental changes, caused by clear-cutting and lim-
perimental plots. In summary, every family of the ing.

Table 6
Population density (ind. m-*) of Coleoptera on the limed (li) and unlimed (unli) clear-cut and the limed (li) and unlimed (unli) beech forest
plots in the Solling mountains (1992/1993), mean density of two plots
Photo-eclectors Funnel method
Clear-cut Forest Clear-cut Forest
li unli li unli li unli ii- unli
CARABIDAE
Noriophilus biguttatus CF.1 1 0 0 0
Trechus quadristriatus (Schrk.) 0 1 7 21
Bembidion lampros (Hbst.) 1 0 0 0
Trichocellusplacidus (Gyllenhal) 0 0 5 +
Pterostichus oblongopunctatus CF.1 9 10 4 2.
Pterostichus metallicus CF.1 0 0 0 0
Trichotichnus laeuicollis (Duft.1 0 0 0 0

HYDROPHILIDAE
Megasternum boletophagum Marsh. 0 0 0 0 0 0 2 0

CATOPIDAE
Nargus wilkini Spence 15 2 7 5 3 4 2 6
Catops longulus Kelln. 0 4 0 0 0 + Cl 0

LIODIDAE
Colenis immunda Sturm 0 0 1 0 0 0 0 0
Agathidium nigripenne F. 0 0 0 1 0 0 + 0

SCYDMAENIDAE
Neuraphes angulatus (MU.) 3 0 1 0 0 + 0 0
Neuraphes sp. 0 0 1 1 0 0 0 0

PI’ILIIDAE
Acrotrichis sp. 5 7 6 2 14 23 119 22

STAI’HYLINIDAE
PROTEININAE
Proteinus brachypterus F. 0 0 1 0 0 0 0 0
Proteinus oualis Er. 1 0 1 0 0 0 0 0

OMALIINAE
Anthophagus angusticollis Mannh. 1 1 2 2 2 0 0 0
Phyllodrepa ioptera (Steph.) 6 4 14 9 0 0 0 0
Dropephylla gracilicornis Fairm. 0 0 0 0 0 + 0 0

OXYTELINAE
Coprophilus striatulus CF.1 1 5 0 1 0 0 0 0

STENINAE
Stenus latifions Er. 4 0 0 0 1 0 0 0

PAEDERINAE
Lathrobium fuluipenne (Grav.) 0 1 0 0 1 0 5 3
Lathrobium brunnipes CF.1 0 0 0 0 1 0 3 0
Lathrobium longulum Grav. 0 0 0 0 + + 0 0
Domene scabricollis (Er.1 1 5 2 1 0 + 11 +
44 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (I 995) 35-51

Table 6 continued
Photo-eclectors Funnel method
Clear-cut Forest Clear-cut Forest
-
Ii unli Ii unli li unli Ii unli

XANTHOLININAE
Xantholinus fricolor (F.) 1 0 0 0 1 I I
Xantholinus linearis (01.) 0 0 0 + 0 0 0
Othiuspunctulatus (Gze.) 1 1 2 3 3 b 3
Othius myrmecophilus Kiesw. 0 1 1 8 IL! 7 37

STAPHYLININAE
Philontus sp. 0 1 0 0 0 0 0 0
Quedius mesomelinus (Marsh.) 1 1 s I 0 0 0 0
Quedius fimatus Steph. 0 0 0 0 0 cl + 0
Quedius xanthopus Er. 0 2 3 2 0 0 0 0
Quedius uexans Epph. 0 1 0 0 0 0 0 0
Quedius maurus (Sahlb.) 0 0 1 0 0 0 0 0
Quediusfulgidus (F.) 0 1 0 0 0 0 0 0
Quedius curtipennis Bernh. 0 0 1 0 0 0 0 0
Quedius sp. 0 1 0 2 0 0 0 0

TACHYPORINAE
Mycetoporus brunneus (Marsh.) 1 0 0 0 0 0 0 0
Bolitobius exoletus Er. 0 0 0 0 0 0 + 0
Tachinus subterraneus CL.) 0 0 0 1 0 0 0 0
Tachinw laticollis (Grav.) 0 0 1 0 0 0 I 0
Tachinus marginellus (F.) 1 0 0 0 0 0 0 0
Tachyporus chrysomelinus L. 0 0 0 0 0 0 0 0

Aleocharinae 9 1s 26 28 117 50 110 78

PSELAPI-IIDAE
Bryaxis nodicornis (AubC) 0 1 0 0 0 0 0 0
Plectophloeus erichsoni (Aubt) 0 1 0 0 0 0 0 0

CANTHARIDAE
Malthodes mysticus Kiesw. 0 0 1 2 0 0 0 0
CantharisFgurata Mann. 1 0 0 0 0 0 0 0
Cantharis obscura L. 0 1 0 0 0 0 0 0
Bhagonycha lignosa (MU) 7 1 1 2 0 0 0 0
Podistra rufotestacea (Letzn.) 0 0 1 0 0 0 0 0
Malthodes sp., females 0 0 0 2 0 0 0 0

ELATEIUDAE
Athous subfkscus F. 3 2 11 19 1 2 3 2
Athous subfkscus, larvae 0 0 0 0 150 168 339 485

BYBRHIDAE
Cytilus sericeus CF.) 0 0 0 0 0 + 0 0

NITIDULIDAE
Brachypterus urticae (F.) 1 0 0 0 0 0 1 0
Thalycra feruida (Olivier) 0 0 0 0 0 0 + 0
Epuraea longula Er. 0 1 0 0 0 0 0 0
Epuraea sp. 0 0 0 0 + 0 0 0
 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51 4s

Table 6 continued
Photo-eclectors Funnel method
Clear-cut Forest Clear-cut Forest
li unli Ii unli li unli Ii unli
RIIIZOPIIAGIDAE
Rhiwphagus dispar (Payk.) 0 2 0 0 0 0 0 0
Rhisophagus ferrugineus (Payk.) 1 3 2 0 0 0 0 0

CRYPTOPHAGIDAE
Atomaria sp. 2 3 3 2 0 0 2 0
Cryptophagus silesiacus Ganglb. 1 0 0 1 0 0 0 0

LATI-IRIDIIDAE
Corticaria sp. 1 16 2 3 0 0 1 0
Cartoderema elongata (Curt.) 0 1 0 0 0 0 0 0
Lathridius nodifer Westw. 8 3 4 0 0 0 0 0

COCCINELLIDAE
Propylaea quatuordecimpunctata CL.1 0 0 0 0 + 0 + 0
Halyzia sedecimguttata (L.) 0 0 0 1 0 0 0 0
Coccinella septempunctata L. 1 0 0 0 0 1 0 1
PY’I’IUDAE
Rhinosimusplanirostris (F.) 0 0 0 0 0 0 0 0
Rabocerus foveolatus (Ljungl 0 0 0 1 0 0 0 0
MORDELLIDAE
Nassipa rufilabris (Gyll.) 0 0 0 0 0 0 0 0
SCARABAEIDAE
Melinopterus prodromus (Brahm) 0 0 1 0 0 0 0 0
CHRYSOMELIDAE
Batophila sp. 0 0 0 0 + 0 0 0
SCOLYTIDAE
Xyloterus domesticus (L.) 0 1 1 1 0 0 0 0
CURCULIONIDAE
Polydrusus undatus (F.) 2 0 6 5 0 0 0 0
Sitona cambricus Steph. 1 0 0 0 0 0 0 0
Strophosoma melanogrammum (F&t.) 1 1 6 14 0 0 1 +
Strophosoma capitatus (Deg.) 0 0 1 2 0 0 0 +
Otiorhynchus singularis CL.) 0 0 1 1 0 0 0 +
Phyllobius argentatus (L.) 0 2 so 70 0 0 0 3
Rhynchaenus fagi (L.) 1 1 0 0 0 0 1 0
Rhinomias forticornis @oh.) 0 0 0 2 0 0 0 1
Ceutorhynchus quadridens Germ. 0 0 0 0 0 0 + 0
Ceutorhynchus erysimi Germ. 1 0 0 0 0 0 0 0
Acalles camelus (F.) 0 0 1 2 0 0 2 1
ASPIDIPIIORIDAE
Aspidiphorus orbiculatus (Gyll.) 1 0 0 0 0 0 0
CLAMBIDAE
Clambus punctulum Beck 1 0 0 0 0 0 0
Methods: ground photo-eclectors (capture period 11 March to 4 November 1992) and funnel extraction method (mean of four sampling
dates 1992). +, between 0 and 1 ind. rn-’ found.
46 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51

By means of artificial trunks, the attraction, which Macroceridae and Keratoplatidae form part of the
tree trunks exert on the soil macrofauna (Chilopoda, zoophages.
Diplopoda, Coleoptera, Dipteral, was examined by Emergence abundance of total zoophagous feed-
Funke and Herlitzius (1984). These authors observed ers was comparable for all experimental plots. With
that Lonchopteridae avoided the artificial trunks. This Empididae and Dolichopodidae, however, a signifi-
finding is consistent with the high number of Lon- cant change became evident: the emergence abun-
chopteridae found on clear-cut plots. dance value for Empididae increased after liming
(P < 0.05; Table 5). Dolichopodid numbers were
3.2.1.4. Zoophages. Rhagionidae, Empididae, Do- significantly higher on the clear-cut plots relative to
lichopodidae, Hybotidae, Pipunculidae, Tachinidae, the beech forest plots (P < 0.01).

Ind lm2
400
a) Lumbricidae

fa Dendiobaena. ~uven~l f7 Dendrobaena octaedra

lnd lm2 !nd /m2

”

::v
c) Curculionidae
4-

HTrechus quadrtstrlatus 0 PI oblongopuncratus

q Trichocellus r?lacldus c]P: metallicus

lnd /m”

80
I 1) Lithobiidae
fi0

-
Cl noI1 Cl /I f0 noll f0 I/

q Aleochmna 001 myrmecophllus

Not punctulatus mother ta L , anamorph q L muiabik q 1 curtlpes bother

Fig. 4. Density of the soil macrofauna (ind. m -2 ) on limed and unlimed clear-cut plots, and on limed and unlimed beech forest plots in the
Soiling mountains, sampled by funnel extraction method (mean of four sampling dates 1992). cl noli, clear-cut, not limed; cl Ii, clear-cut,
limed; fo noli, beech forest, not limed; fo li, beech forest, limed.
 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51 47

3.3. Coleoptera and removing of the trees in autumn 1989, patches of

the upper strata of the soil, the preferred habitat of
A total of 24 Coleoptera taxa were recorded, with Athous subfuscus larvae (Moritz, 1986), were af-
86 species/groups identified (Table 6). In funnel fected (J. Schauermann, personal communication,
samples Aleocharinae were most abundant, averag- i993). During this time, the larvae, which could be
ing 89 ind. mV2. They were followed by Acrotrichis expected to hatch in the year of the present study,
spp. (44 ind. rnm2> and Othius myrmecophilus (12 were already present in the soil. It could be con-
ind. me2). In the catches with the ground photo- cluded that the decrease in Athous subfuscus after
eclectors Phyllobius argentatus (30 ind. mm2) was clear-cutting was caused by the death of the larvae
most abundant, followed by Aleocharinae (19 ind. due to the experimental treatment.
mp2) and Athous subfuscus (9 ind. mw2) (Table 6).
The average population density of Coleoptera was
3.3.2. Curculionidae
175 ind. rnp2 (funnel method) and 224 ind. me2
Eight curculionid species were detected. In the
(ground photo-eclectors) (Fig. 3~). By means of the
funnel catches Phyllobius argentatus prevailed com-
funnel method, on the clear-cut plots, an average of
prising 32% of the total Curculionidae, followed by
130 ind. rnp2 was found, whereas comparatively
Acalles camelus (26%) and Strophosoma melano-
high average numbers were maintained on the beech
grammum (12%). Clear-cutting resulted in a com-
forest plots (219 ind. m-2 ). The density of Coleoptera
plete breakdown of the curculionid population. On
was higher on the limed plots (217 ind. me2) than
the beech forest plots the average abundance of
on the unlimed plots (132 ind. me2 ) (funnel method)
Curculionidae was 5 ind. rne2. No specimen of
(Fig. 3(c)). By means of the ground photo-eclectors,
Curculionidae was found on the clear-cut plots (Fig.
on the clear-cut plots, an average of 92 ind. rnw2
4(c) and Table 6). This agrees with the results of the
was found and 179 ind. m-‘, on average, on the
eclectors, where the curculionid population was re-
beech forest plots (Fig. 3(c)). Reasons for these
duced on the clear-cut plots, with a significance level
differences are discussed in the following chapters
of P < 0.001 (Fig. 3(d)). While comparatively high
dealing with the dominating families.
numbers were maintained on the beech forest plots
Perel (1965) found a decrease in the total density
(80 ind. me2 on average), on the clear-cut plots only
of Coleoptera after cutting, whereas Huhta et al.
5 ind. me2 emerged on average. The effect of
(1967) reported an increase in Coleoptera soon after
clear-cutting on Polydrusus undatus (P < 0.051,
clear-cutting.
Strophosoma melanogrammum (P < 0.001) and
Phyllobius argentatus (P < 0.001) was a significant
3.3.1. Larvae of Athous subfuscus (Elateridae)
decrease in emergence abundance.
On the beech forest plots, the mean abundance of
How can the strong decline in Curculionidae on
the larvae of Athous subfiscus was 412 ind. rne2,
the clear-cut plots be explained? The adults of most
whereas comparatively low numbers were main-
of the Curculionidae found in this study feed on
tained on the clear-cut plots (Fig. 4(b) and Table 6).
leaves in the crowns of beech trees. After they
Correspondingly, the emergence abundance of
emerge from the soil, the adult beetles either fly into
Athous subfuscus was significantly (P < 0.001) re-
the crowns of the beech trees or they climb up the
duced on the clear-cut plots.
tree trunks. It makes sense, therefore, that the fe-
Funke and Herlitzius (1984) observed that Elateri-
males choose oviposition sites close to beech trees.
dae were attracted by artificial tree trunks. These
Funke and Herlitzius (1984) reported on the attrac-
observations agree with the decline in Athous sub-
tion of beech trees for the adults of the Curculion-
j&us population after clear-cutting.
idae Polydrusus sp. and Strophosoma sp.
Since the larvae of Athous subfuscus stay in the
soil for 3-5 years (Moritz, 1986) and since clear-cut-
ting was performed 3 years ago, a direct influence of 3.3.3. Carabidae
the experimental treatments on the larvae of Athous Seven species of Carabidae were found. Of these,
subfuscus must be considered. During the cutting Trechus quadristriatus, Pterostichus oblongopuncta-
48 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51

tus, Trichocellus placidus and Pterostichus metalli-
cus were most abundant (Table 6).
On the limed plots the average density of Cara-
bidae was higher (20 ind. m-* by the funnel method
and 12 ind. mm2 using ground photo-eclectors) than
on the unlimed plots (3 and 11 ind. m-‘, respec-
tively) (Fig. 4(d); funnel method only). Similar pat-
terns were observed for the individual species. Of
Trechus quadristriatus on the limed plots, an aver-
age of 14 and 2 ind. rnm2, respectively, were found,
while on the unlimed plots an average of only 1 and
< 1 ind. m-‘, respectively, were found. The abun-
dance of Trichocellus placidus was highest on the
limed clear-cut plots (an average of 3 and < 1 ind.
rnb2, respectively). In the catches with the funnel
method, liming had been slightly favourable for
Pterostichus oblongopunctatus. Furthermore, clear-
cutting had a positive effect on the population den-
sity of Pterostichus oblongopunctatus. The average
density of this species was 3 ind. rn-’ on the
clear-cut plots and 1 ind. mm2 on the beech forest
plots. No tendency was found for this species in the
samples taken with the ground photo-eclectors.
Knie (1975) reported that the density and species
variety of the carabid fauna in the humid and acidic
forests near Bonn (Germany) depend on the amount
of vegetation. He found that a low density of the
herb layer caused a decline in species numbers and
an increase in population density. In clear-cuts, where
the herb layer was well developed, Knie detected the
highest number of species, but only a small popula-
tion density. In the present study, however, no obvi-
ous change in species numbers was found. Further-
more, the population density was highest on the plots
with a dense herb layer (limed plots).
ground photo-eclectors, whereas the funnel method
revealed a decline in the population density of
Staphylinidae after clear-cutting (P < 0.01) (Fig.
4(e)). The same response was observed in Aleochari-
nae (P < 0.051. Furthermore, the liming treatment
caused an increase in the abundance of Aleocharinae
(P < 0.05). Othius punctulatus presented no evident
pattern. On the unlimed beech forest plots, the aver-
age density of Othius myrmecophilus was five times
as high (37 ind. m-*) as on the limed beech forest
plots (7 ind. rne2). Clear-cutting had a negative
effect on the population of Lathrobium fklvipenne.
No individual of this species was detected on the
unlimed clear-cut plots. On the limed clear-cut plots,
the abundance of this species was relatively low. On
the beech forest plots, however, an average of 4 ind.
me2 of Lathrobium jklvipenne were collected. For
Lathrobium brunnipes similar patterns were ob-
served (Table 61.
Huhta et al. (19671 found an increase in
Staphylinidae soon after clear-cutting, while Szujecki
(1971, 1972) reported a decline in Staphylinidae in
clear-cut areas.
3.4. Lithobiidae
The average density of Lithobiidae was found to
~JJ lxveemu 31 ind. me2 (unlimed forest plots) and
. -2 (limed forest plots) (Table 7). Four
species were identified, the predominant one being
Lithobius mutabilis, averaging 18 ind. rn--‘. The
density of Lithobius curtipes was 15 ind. mm-‘. Just
one individual of Lithobius macilentus and one of
Lithobius dentatus were found.

Table I
Population density (ind. mm2 ) of Lithobiws spp. on the limed (Ii)
3.3.4. Staphylinidae and unlimed (unli) clear-cut and the limed (Ii) and unlimed &mli)
The average abundance of Staphylinidae was 114 beech forest plots in the Solliig motintains (1992/1!B3)
ind. me2 (funnel method) (Fig. 4(e)) and 38 ind. Clear-cut Forest -
m-2 (ground photo-eclectors) (not shown). In the
ii----- unli Ii unli
catches using the funnel method, 78% of the
Lithobius curt@ C.L. Koch 17 10 24 9
Staphylinidae were Aleocharinae, 14% were Othius
Lithobius mutabilis L. Koch 14 19 28 I2
myrmecophilus and 3% were Othius punctulatus. In Lithobius macilentus L. Koch 0 + 0 0
the catches with ground photo-eclectors, 51% of the Lithobiusdentutus C.L. Koch 0 0 i 0
Staphylinidae were Aleocharinae and 17% Phyllo- Lithobius, juvenile 19 13 28 10
drepa ioptera. No obvious differences between the Funnel method (mean of four sampling dates 1992). + , between 0
treatments were observed in the samples taken with and 1 ind. me2 found.
 A. Theenhaus, M. Schaefer/Forest
The average density of all Lithobius spp. was
higher on the limed plots (65 ind. m-‘) than on the
unlimed plots (37 ind. m-‘1 (P < 0.001) (Fig. 4(f)).
The same was true for the anamorphic Lithobiidae,
with a significance of P < 0.01. Lithobius curtipes
thrived on the limed plots as well. An increase in
Lithobius numbers, due to liming, was described by
Schauermann (1985). Schaefer (1991~) stated that
Centipedes are an important group of the predatory
macrofauna in the soil-litter subsystem. Lithobiidae
feed mainly on mobile animals such as Collembola,
Diptera and Lumbricidae. It may therefore be as-
sumed that the relatively high lumbricid numbers on
the limed plots have contributed to the increase in
Lithobius.
Gretschy (1952) pointed out that Myriopoda need
highly porous soil because they lack digging ability.
It is possible that the high number of Lumbricidae on
the limed plots increases the porosity of the soil, thus
improving the microhabitat for the Lithobiidae.
4. General conclusions
In this study, the change of soil moisture, caused
by the treatments, was assumed to be the most
important factor determining the change in popula-
tion densities. Other factors, such as soil and litter
chemical parameters, were not considered because of
the speculative nature of the influence of these fac-
tors on the abundance of faunal groups.
It has been hypothesized that liming changes
moder profile into mull profile. Schaefer and
Schauermann (1990) compared the soil fauna charac-
teristics of two German beech forests-one on mull
soil, the other on moder soil (Soiling). They found
that Lumbricidae, Chilopoda and carabid beetles are
favoured on mull soil. The results of the present
study indicate a positive effect of liming on the
abundance of these taxa and, therefore, a change
from moder into mull soil. Mull soils are character-
ized by a much higher total soil-fauna biomass than
moder and mor soils (Schaefer, 1991~). This is be-
cause mull soils are dominated by Lumbricidae.
Schauermann (1986) points out that consumption
Ecology and Management 77 (1995) 35-51 49
rates by saprophages are lower in the forest on
moder soil (Solling beech forest) than in the beech
mull soil, presumably because mean annual biomass
of all saprophages is only 5-6 g rnp2 in the Solling
forest. In the present study Lumbricidae and other
saprophages such as Sciaridae, Scatopsidae and Tip-
ulidae were most abundant on the limed plots. These
groups contribute significantly to leaf-litter break-
down by their feeding activities. Altmiiller (1976)
pointed out that Diptera are the most important
decomposers in the beech forest of the Solling. The
author concluded that the decomposition of litter can
be attributed mainly to the activity of sciarid and
sciophilid larvae. According to Altmiiller (19761, the
larvae of the Sciaridae and Mycetophilidae (mainly
Sciophilidae) are consuming 13-29% of the annual
litter fall. Hovemeyer (in Schaefer, 1982) estimated
that larvae of Sciaridae in a German mull forest
processed about 8% of the annual input of litter
originating from green phytomass. Binns (1981) as-
sumed that macrohumiphagous Sciaridae and Scio-
philidae have a positive effect on the development of
microflora. Perel et al. (1971) found a stimulating
effect of tipulid larvae (as macrohumiphages) on the
decomposition process. Summing up, the results of
the present study suggest a change of moder soil into
mull soil after liming.
Heterotrophic effects are greatest in decomposer
processes. In a forest ecosystem about 90-95% of
total heterotrophic respiration is contributed by de-
composers. Furthermore, the heterotrophic commu-
nity performs an important function in controlling
the fluctuations in the rate of nitrogen release
(Schaefer, 1991b). Therefore, because of the in-
creased density of Lumbricidae, Sciaridae, Scatopsi-
dae and Tipulidae on the limed plots, a more rapid
mineralization and mobilization of nutrients was ex-
pected. This, however, was not found. Instead, on
the limed plots (19911, less nitrogen was mineralized
compared to the unlimed plots.
It has further been hypothesized that the decom-
poser community decreases as a result of clear-cut-
ting. This, however, has not been found. Lumbricids
were not significantly influenced by clear-cutting.
This was also true for the macrohumiphagous
Diptera. The emergence abundance of Limoniidae on
the clear-cut plots was found to be even significantly
higher than on the forest plots.
50 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-5 1

Acknowledgements Hennig, W. (Editor), 1968. Die Larvenformen der Dipteren.

Akademie, Berlin, 1271 pp.
We thank Dr. J. Schauermann for helpful assis- Heynen, C., 1988. Zur Biologie eines Buchenwaldbodens. 1 I. Die
Dipterenlarven. Carolinea, 46: 115-130.
tance and many fruitful discussions. For advice on
Hovemeyer, K., 1984. Die Dipterengemeinschaft eines Buchen-
statistical matters we are indebted to Dr. Michael waldes auf Kalkgestein: Produktion an Imagines, Abundanz
Judas. The study was supported by the Bundesminis- und rliumliche Verteilung inbesondere der Larven. Pedobiolo-
terium fir Forschung und Technologie. gia, 26: l-15.
Hovemeyer, K., 1985. Die Zweifliigler (Dipteral eines Kalk-
buchenwaldes: Lcbenszyklen, Raum-Zeit-Muster und Nah-
References rungsbiologie. Thesis, Gottingen, 280 pp.
Hovemeyer. K., 1991. The study of dipterous populations and
Altmiiller, R., 1976. Zum Energieumsatz von Dipterenpopulatio- communities in European terretrial ecosytems. In: L. Weis-
nen im Buchenwald (Luzulo-Fagetum). Thesis, Glittingen. mann, 1. Orszagh and A. C. Pont (Editors), Proceedings of the
Altmiiller, R., 1979. Untersuchungen liber den Energieumsatz von Second International Congress of Dipterology. SPB Academic
Dipterenpopulationen im Buchenwald (Luzulo-Fagetum). Pe- Publishing, The Hague, pp. 99-109.
dobiologia, 19: 245-278. Hovemeyer. K., 1992. Die Dipterengemeinschaft eines Kalk
Bauhus, J., 1994. Stoffumsatze in Lochhieben. Berichte des buchenwaldes: Eine siebenjlhrige Untersuchung. Zool. Jb.
Forschungszentrums Waldlikosysteme, A 113. Thesis, 119: 225-260.
Gdttingen. Huhta, V.. 1976. Effects of clear-cutting on numbers, biomass and
Bauhus, J. and Barthel, R., 1994. Mechanisms for carbon and community respiration of soil invertebrates. Ann. Zool. Fenn..
nutrient release and retention in beech forest gaps. II. The role 13: 63-80.
of soil microbial biomass. Plant Soil, 37: 17-24. Huhta, V., 1979. Effects of liming and deciduous litter on earth-
Bauhus, J. and Bartsch, N., 1995. Mechanisms for carbon and worm (Lumbricidael populations of a spruce forest, with an
nutrient release and retention in beech forest gaps. I. Microcli- inoculation experiment on Allolobophoru caliginosa. Pedohi-
mate, water balance and seepage water chemistry. Plant Soil, ologia, 19: 340-345.
168-169: 579-584. Huhta, V., Karppinen, E., Nurminen, M. and Valpas, A., IY67.
Binns, ES., 1981. Fungus gnats (Diptera: Mycetophilidae/ Effect of silvicultural practices upon arthropod, annelid and
Sciaridae) and the role of mycetophagy in soil: a review. Rev. nematode populations in coniferous forest soil. Ann. Zoo1
Ecol. Biol. Sol., 18: 77-90. Fenn.. 4: 87-143.
Brauns, A. (Editor), 1954. Terricole Dipterenlarven. Giittingen, Irmler, U. and Heydemann, B., lY89. Der Einfluss einer Kalk-
Musterschmidt, 179 pp. mergelbehandlung auf die Bodenfauna dreier schleswig-hol-
Cramer, E., 1968. Die Tipuliden des Naturschutzparkes Hoher steinischer Waldtypen. Verh. Ges. Gkol., 17: 591-596.
Vogelsberg. Dtsch. Entomol. Z., N.F. 15(1/3): 133-231. Jacobs, W. and Renner, M. (Editors), 1988. Biologie und Gkolo-
Ellenberg, H. (Editor), 1971. Integrated experimental ecology. gie der Insekten, 2. ilberarbeitete Auflage. Fischer, Stuttgart,
Methods and results of ecosystem research in the German 690 PP.
Soiling project. Ecol. Stud., 2: 1-214. Kempson, D., Lloyd, M. and Gherardij, R., 1963. A new extractor
Ellenberg, H., Mayer, R. and Schauermann, J. (Editors), 1986. for woodland litter. Pedobiologia, 13: l-21.
Gkosystemforschung, Ergebnisse des Sollingprojektes. Ulmer, Knie, J.. 1975. Vergleichend-Gkologische Untersuchungen der
Stuttgart, 507 pp. Carabidenfauna verschiedener Standorte des Kottenforstes her
Feldmann, R., 1992. Die Bodenmakrofauna im Lennebergwald. 1. Bonn. Decheniana, 128: 3-19.
Die Dipteren. Mainz. Naturwiss. Arch., 30: 171-241. Moritz, G., 1986. Zur Gkologie von Schellkafem (Coleoptera:
Funke, W., 1971. Food and energy turnover of leafeating insects Elateridael in Buchen- und Fichtenwlildem. Thesis, Clottingen.
and their influence on primary production. Ecol. Stud., 2: Odum, E.P., 1971. Fundamentals of Ecology, 3rd edn. Saunders
81-93. Philadelphia, 574 pp.
Funke, W., 1991. Tiergesellschaften in Waldern. Kern- Okland. B., 1994. Mycetophilidae (Dipteral, an insect group
forschungszentrum, Karlsruhe. vulnerable to forestry practices? A comparison of clear-cut.
Funke, W. and Herlitzius, H., 1984. Zur Orientierung von Arthro- managed and semi-natural spruce forests in southern Norway.
poden der BodenoberfHche nach Stammsilhouetten im Wald. Biodiversity Conserv., 3: 68-85.
Jahresber. Naturwiss. Ber. Wuppertal, 37: 8-13. Perel, T.S.. 1965. Die Bodenbewohner der natnrlichen Fich-
Gretschy, G., 1952. Die Sukzession der Bodentiere auf Fichten- tenbestande in der siidlichen Taiga und ihre Ver%ulerungen im
schllgen. Veroeff. Bundesanst. Alp. Landwirtsch. Admont, 6: Zusammenhang mit Kahlschlag und Holzartenwechsel. Pedo
25-85. biologia, 5: 102-121.
Healey, I.N. and Russel-Smith, A., 1971. Abundance and feeding Perel, T.S., Karpachevsky, L.O. and Yegorova, E.V., 1971. The
preference of tly larvae in two woodland soils. 4. Colloq. role of Tipulidae (Dipteral larvae in decomposition of forest
Pedobiol. Dijon, 1970. Ann. Zool. Ecol. Anim. Numdro hors- litterfall. Pedobiologia, 11: 66-70.
serie, Paris 3: 177-191. Piearce, T.G.. 1972. Acid intolerant and ubiquitous Lumbricidae
 A. Theenhaus, M. Schaefer/Forest Ecology and Management 77 (1995) 35-51 51

in selected habitats in North Wales. J. Anim. Ecol., 41: Schauermann, J., 1982. Verbesserte Extraktion der terrestrischen
397-410. Bodenfauna im Vielfachgemt modifiziert nach Kempson und
Raw, F., 1967. Arthropoda (except Atari and Collembola). In: A. MacFadyen. Kurzmitteilungen aus dem Sonderforschungsbe-
Burges and F. Raw (Editors), Soil Biology. Academic Press reich, 135: 47-50.
Inc., London, pp. 323-362. Schauermann, J., 1985. Zur Reaktion von Bodentieren nach
Schaefer, M., 1982. Zur Funktion der saprophagen Bodentiere Dbngung von Hainsimsen-Buchenw%ldern und Siebenstem-
eines Kalkbuchenwaldes: Ein langfristiges Untersuchungspro- Fichtenforsten im Solling. Allgem. Forstzeitschr., 43: 1159-
gramm im Gottinger Wald. Drosera, 82: 75-82. 1160.
Schaefer, M., 1991a. The animal community: diversity and re- Schauermann, J., 1986. Die Tierwelt, ihre Nahrungsbeziehungen
sources. In: E. Rijhrig and M. Uhich (Editors), Temperate und ihre Rolle. Siedlungsdichten und Biomasse. In: H. Ellen-
Deciduous Forests (Ecosystems of the World). Elsevier, Am- berg, R. Mayer and J. Schauermann (Editors), Gkosystem-
sterdam, pp. 51-120. forschung, Ergebnisse des Sollingprojektes. Ulmer, Stuttgart,
Schaefer, M., 1991b. Secondary production and decomposition. pp. 225-265.
In: E. Riihrig and M. Ulrich (Editors), Temperate Deciduous Swift, M.J., Heal, O.W. and Anderson, J.M., 1979. Decomposi-
Forests (Ecosystems of the World). Elsevier, Amsterdam, pp. tion in Terrestrial Ecosystems. Blackwell, Oxford, 372 pp.
175-218. Szujecki, A., 1971. The effect of clear-cutting upon the commu-
Schaefer, M., 1991~. Fauna of the European temperate deciduous nity of litter inhabiting Staphylinids (Co]., Staphylinidae) in
forest. In: E. Rohrig and M. Ulrich (Editors), Temperate fresh pine forests. Folia For. Pol. Ser. A 18: 5-45.
Deciduous Forests (Ecosystems of the World). Elsevier, Am- Szujecki, 1972. Impact of clear-cutting on the soil entomofauna.
sterdam, pp. 503-526. Contribution in the 7th World Forestry Congress, Buenos
Schaefer, M. and Schauermann, J., 1990. The soil fauna of beech Aires 1972.
forests: comparison between a mull and a moder soil. Pedobi-
ologia, 34: 299-314.