PERGAMON Soil Biology and Biochemistry 31 (1999) 517±523

Earthworm and soil moisture e€ects on the productivity and

structure of grassland communities
J.G. Zaller *, J.A. Arnone III 1
Botanisches Institut, UniversitaÈt Basel, SchoÈnbeinstrasse 6, 4056 Basel, Switzerland
Accepted 9 July 1998

Abstract

The objectives of this study were (1) to evaluate the e€ect of earthworm activity on aboveground plant biomass production of
native calcareous grassland communities in NW-Switzerland and (2) to determine which plant functional types (graminoids,
non-legume forbs and legumes) are most responsive as indicators of potential e€ects on plant community structure. Earthworm
activity was manipulated in the ®eld by creating three earthworm densities (low: 37, ambient: 114, high: 169 worms m ÿ 2) and
two soil moisture conditions (ambient and 280 mm yr ÿ 1 additional rain) in 30 1  1 m2 trenched plots (to a depth of 45 cm with
nylon screening). Earthworm density was censused and readjusted in the spring and autumn of 1996 and again in the spring of
1997 using the Octet electro-sampling method. Earthworm activity, measured as cumulative surface cast production, was
signi®cantly di€erent among worm density treatments (low worm density: 591 2 49, ambient: 9912 87, high: 14692 120 g cast
d.m. m ÿ 2 yr ÿ 1), P < 0.001. Cast production increased with increasing worm density, however increased soil moisture, which
was signi®cantly higher in plots receiving additional rain, did not a€ect worm activity at any worm density level. Surprisingly,
earthworm activity had no e€ect on the aboveground biomass production of the plant community or of any plant functional
type; however additional rain stimulated aboveground biomass production of graminoids (+30%, P = 0.006) and Carex spp.
(+200%, P = 0.020). The lack of a stimulatory e€ect of increased earthworm activity on plant biomass production indicates
that apparent earthworm-induced increases in plant nutrient availability were insucient to promote the growth of the perennial
plant species in these native grasslands. Our results, taken together with those from earlier studies in these grasslands, suggest
that the e€ects of earthworm activity on plant community structure occur slowly where they relieve soil constraints on plant
growth. # 1999 Elsevier Science Ltd. All rights reserved.

1. Introduction (e.g. Stockdill, 1982; Hoogerkamp et al., 1983; Curry

The activity of earthworms has been shown to
enhance plant yield (Waters, 1951; Sears and Evans,
1953; Stockdill, 1959; Barley, 1961; Stockdill and
Cossens, 1966; Watkin and Wheeler, 1966; Mackay et
al., 1982; Stockdill, 1982; Hoogerkamp et al., 1983;
Curry and Boyle, 1987) because of their bene®cial
e€ects on the soil environment (e.g. Lee, 1985;
Edwards and Bohlen, 1996). However, these results are
derived solely from experiments where earthworms
were introduced to intensively managed grasslands
* Corresponding author. Present address: Research Institute of
Organic Agriculture (FiBL), Ackerstrasse, 5070 Frick, Switzerland.
Tel.: +41-62-865-7243; Fax: +41-62-865-7273; E-mail: hans.zaller
@®bl.ch.
1
Present address: Desert Research Institute, Biological Sciences
Center, P.O. Box 60220, Reno, NV 8956, USA.
and Boyle, 1987) or where the e€ects of earthworm ac-
tivity on plant growth may be confounded with the
other e€ects of the land management practice that
resulted in a particular earthworm density (e.g.
Kladivko and Timmenga, 1990; Edwards et al., 1995).
Thus, the extent to which these ®ndings may be
applied to extensively managed or natural grasslands is
questionable.
Earthworms and their activity may also a€ect the
structure of plant communities by in¯uencing the con-
tribution of various plant species to the community
(e.g. Hopp and Slater, 1948; van Rhee, 1965;
Hoogerkamp et al., 1983; Thompson et al., 1993;
Zaller and Arnone, 1998). This may occur via selective
seed burial and dispersion (Thompson et al., 1994;
Willems and Huijsmans, 1994) or by promoting the
establishment and growth of particular plant species as
a result of a plant's proximity to nutrient-rich casts

0038-0717/99/$19.00 # 1999 Elsevier Science Ltd. All rights reserved.

PII: S 0 0 3 8 - 0 7 1 7 ( 9 8 ) 0 0 1 2 6 - 6
518 J.G. Zaller, J.A. Arnone III / Soil Biology and Biochemistry 31 (1999) 517±523
(Zaller and Arnone, 1998) and the ability of a plant
species to exploit these nutrients (Hofer, unpublished
data).
The objectives of our study were: (1) to evaluate the
e€ects of earthworm activity on aboveground biomass
production of native calcareous grassland communities
and (2) to determine which plant functional types (gra-
minoids, nonleguminous forbs and legumes) are most
responsive as an indication of potential e€ects on
plant community structure. To do this, earthworm ac-
tivity was manipulated in the ®eld for two growing
seasons: ®rstly, by manipulating the size of earthworm
community in trenched ®eld plots and, secondly, by
increasing soil moisture in these plots.
2. Materials and methods
2.1. Site description
The calcareous grassland we studied is located on a
208 southwest-facing slope near the village of
Nenzlingen (canton Basel-Land), NW-Switzerland (500
m a.s.l., 47827 0 N, 7834 0 E). Mean annual precipitation
is about 920 mm at mean air temperatures of about
8.58C (Ogermann et al., 1994). Up to 1993 this grass-
land had been used for extensive cattle grazing and
since 1993 the area has been fenced and mown twice a
year in spring and autumn. Among the 100 vascular
plant species found on this site, the grass Bromus erec-
tus is dominant (Zoller, 1954; Huovinen-Hufschmid
and KoÈrner, 1998). Soils are classi®ed as a transition
Rendzina (pH is about 6.5, bulk density of the top soil
1.1 g cm ÿ 3, C-to-N ratio about 12), with a well devel-
oped, stone-free, loamy topsoil and a rapid transition
at 15±25 cm depth to the underlying calcareous scree
material (Ogermann et al., 1994).
2.2. Experimental design
To control earthworm density, 30 1  1 m plots were
trenched to a depth of 45 cm with 1-mm-mesh nylon
window screen in spring 1995. The screen extended 15
cm above the soil surface to create an aboveground
earthworm barrier. Trenching to 45 cm in these shal-
low soils would be expected to strongly limit subsur-
face lateral movement of earthworms into or out of
the plots. Plots were arranged in a randomized com-
plete block design (5 blocks), with three earthworm
densities (low, ambient and high) and two amounts of
rainfall (ambient and 280 mm yr ÿ 1 additional rain).
These amounts of added rain were applied to the
appropriate plots during dry periods in the growing
season to maintain sucient soil moisture for earth-
worms to stay active in all but the driest periods.
Volumetric soil water content was continuously moni-
tored over the topmost 10 cm of the topsoil using
time-domain-re¯ectrometry (one measurement every 20
min). We also continuously recorded soil temperature
in each plot with thermistors placed at depths of 5 and
15 cm (one reading h ÿ 1).
Earthworm density treatments were established in
May 1996 by ®rst extracting the earthworms from
each plot using the `Octet' electro-sampling method
(Thielemann, 1986). This non-destructive method has
been shown to provide comparable estimates of earth-
worm community size (Vetter, 1996) and composition
(Zaller and Arnone, 1998) to other more conventional
sampling methods, as long as earthworms are sampled
at times when they are active and when soil moisture
is sucient. To sample, eight steel rods (65 cm
long  6 mm diameter) were inserted into the soil at
equal distances along the circumference of a 60-cm-di-
ameter circle in the center of each plot. An electrical
voltage (300±500 V) was applied to the moist soil over
the entire length of the rods in 4 s pulses supplied
sequentially to opposing pairs of rods in the `Octet'.
The portable transformer (DEKA 4000, Deka
GeraÈtebau, Germany) providing the electrical pulses
was powered by a 12 V car battery. All earthworms
arriving at the soil surface within 40 min in each plot
were collected and stored in cool water until they were
identi®ed, counted and weighed. A total of 9 earth-
worm species were collected (nomenclature follows
BoucheÂ, 1972) representing three ecological groups
(BoucheÂ, 1977): anecics (Nicodrilus longus Ude., N.
nocturnus Ev., Lumbricus terrestris L.), endogeics (N.
caliginosus Sav., Allolobophora chlorotica Sav., A. rosea
Sav., Octolasion cyaneum Sav.) and epigeics (L. casta-
neus Sav., Dendrobaena mammalis Ger.).
All of the earthworms in each ecological group col-
lected from the six plots in each block were pooled;
one-third were added to the ambient-density plots and
the remaining two-thirds to the high-density plots.
This procedure was repeated for each of the ®ve blocks
in May 1996 (start of density treatment) and again in
September 1996 and May 1997 to maintain density
treatments.
Aboveground plant biomass was mowed to a height
of 5 cm in each plot several days before each earth-
worm sampling date in three steps. First, the 10-cm-
wide strip on the perimeter of each plot was removed
and discarded. Second, all the material in an 80  80
cm centre area in each plot, with the exception of two
25  25 cm areas, was harvested, dried at 808C and
weighed. Third, the biomass on the two 25  25 cm
areas in each plot was harvested and sorted into func-
tional groups [graminoids without Carex spp., Carex
spp. (Carex ¯acca and C. caryophyllea), nonlegume
forbs and legumes], dried at 808C and weighed. The
total amount of plant biomass harvested in May and
September 1997 from the centre area in each plot,
 J.G. Zaller, J.A. Arnone III / Soil Biology and Biochemistry 31 (1999) 517±523 519

including that from the two small sub-plots, was used Systat, Evanston, IL; Wilkinson et al., 1992). All error
to estimate the 1997 aboveground biomass production estimates given in the text are standard errors of the
of the plant community. The two sub-plots were used mean (2S.E.).
to estimate the biomass responses of plant functional
types.
Cumulative surface cast production (dry mass) was 3. Results
measured biweekly from October 1996 to May 1997
on one of the two 25  25 cm sub-plots in each plot as Density and biomass of earthworms as well as earth-
an indicator of earthworm activity. Cast production worm community structure in the 30 plots were similar
was measured only during this period because this is before treatments were imposed (density for anecics,
when earthworm activity is highest (see Zaller and endogeics and epigeics was 672 1, 66 27 and 12 2 1
Arnone, 1997) and because casts could be seen and worms m ÿ 2, respectively; biomass for anecics, endo-
quanti®ed most reliably during this period since plant geics and epigeics was 62 22, 53 25 and 32 0.5 g
biomass was lowest and because we needed to avoid m ÿ 2, respectively). The manipulation of earthworm
disturbing the plant canopy during the growing season. community size (density and biomass) was indeed
After weighing (fresh mass) each cast in the ®eld on a e€ective (Fig. 1), although earthworm populations
portable balance, it was returned to its original pos- ¯uctuated between sampling dates with community
ition and deformed slightly to facilitate the identi®- size (number and biomass) in low-density plots tending
cation of newly produced casts at the next sampling to increase and in high density plots tending to
date. Cast subsamples from each plot were taken at decrease, whereas in ambient density plots remaining
each sampling date to calculate fresh mass-to-dry mass largely unchanged. Low density plots contained fewest
ratios (808C, 24 h). earthworms (37 25 worms m ÿ 2) and least biomass
(26.72 3.3 g m ÿ 2) on average; the ambient density
2.3. Statistical analysis plots contained about twice that and the high density
plots about 50% more than the ambient plots con-
The mean density and biomass of the earthworm
community in each plot over the course of the exper-
iment was calculated based on ®ve sets of data: the
May 1996 establishment data, the September 1996
electro-sampling and re-establishment data and the
May 1997 electro-sampling and re-establishment data.
The e€ect of the worm density treatment on actual
worm density and biomass was tested using the mean
earthworm density (and biomass) for each plot in an
analysis of variance (ANOVA) with earthworm density
(`worm'), `rain' and `block' as factors. The e€ect of the
rain treatment on soil moisture was tested using the
mean daily volumetric soil water content from May
1996 to September 1997 for each plot in an ANOVA
with the same three factors. The e€ects worm density
and rain treatments on earthworm activity were tested
using total cumulative surface cast production from
October 1996 to May 1997 in each plot using the same
ANOVA procedure.
Worm density and rain e€ects on aboveground bio-
mass production of the plant community and each
functional group, as well as that of Carex spp. pro-
duction was similarly tested by ANOVA. We used
simple linear regression to assess relationships between
the 1997 aboveground biomass production of the plant
community (dependent variable) and mean earthworm
community density, biomass and activity (independent
Fig. 1. Mean earthworm density (a) and earthworm activity
variables). ANOVAs were used to test the signi®cance
measured as cumulative surface cast production (b) in ®eld plots
of the regression coecient of these linear models with manipulated earthworm density (low, ambient, and high den-
(Sokal and Rohlf, 1981). All statistical analyses were sity) receiving no additional rain and additional rain (mean2S.E.,
performed using Systat (SYSTAT, version 5.2.1, n = 5).
520 J.G. Zaller, J.A. Arnone III / Soil Biology and Biochemistry 31 (1999) 517±523

tained. However, additional rain had no apparent
e€ect on the size of earthworm communities in any
worm density treatment (F = 1.14, P = 0.299; Fig. 1a).
As expected, increasing worm density treatment also
resulted in signi®cant increases in earthworm activity
(cumulative surface cast production; F = 24.27,
P < 0.001; Fig. 1b). Surprisingly, however, additional
rain had no signi®cant e€ect on earthworm activity in
any worm density treatment (F < 0.01, P = 0.976;
Fig. 1b) even though soil water content was consist-
ently greater in these plots (Fig. 2a). The daily mean
soil temperature in all plots ¯uctuated in a normal
fashion with season (Fig. 2b), but did not di€er signi®-
cantly among worm density or rain treatments, or
between 5 and 15-cm depths, at any time during the
study.
Aboveground biomass production (>5 cm height)
of plant communities in 1997 averaged about 440 g
m ÿ 2 in plots receiving no additional rain and 580 g
m ÿ 2 in plots receiving additional rain (F = 24.20,
P < 0.001; Fig. 3a). However, earthworm density
had no e€ect on plant biomass production in either
rain treatment (F = 0.45, P = 0.647; Fig. 3a).
Consequently, biomass production was not correlated
with either earthworm activity (Fig. 3b), actual mean
earthworm density or actual mean earthworm biomass
(data not shown). Additional rain stimulated above-
ground biomass production of graminoids (+30%;
F = 9.30, P = 0.006) and of Carex species (+200%;
F = 6.44, P = 0.020), however worm density treat-
ments had no e€ect on biomass production of these
plant functional groups (Fig. 4). Non-legume forbs
and legumes did not respond to either treatment.
4. Discussion
The absence of any apparent e€ects of earthworm
activity on either the aboveground production of plant
communities (Fig. 3) or individual plant functional
types (Fig. 4) in this calcareous grassland (after 2 yr of
manipulation of earthworm densities) was somewhat
surprising in light of the relatively large stimulatory
e€ects observed in intensively managed grasslands (e.g.
Sears and Evans, 1953; Stockdill and Cossens, 1966;
Curry and Boyle, 1987) and in microcosm studies (e.g.
Hopp and Slater, 1948; van Rhee, 1965). The lack of a
worm e€ect in our study was also puzzling because
earthworm activity, and presumably nutrient avail-
ability to plants (e.g. Scheu, 1994; Willems et al., 1996;
Blair et al., 1997; Zaller and Arnone, 1997), did vary

Fig. 2. Seasonal ¯uctuations in the mean daily volumetric soil water content of the top 10 cm (a) and the mean soil temperature at 5-cm depth
(b) in ®eld plots with manipulated earthworm density receiving no additional rain (lower curve in (a)) and additional rain (upper curve in (a)).
 J.G. Zaller, J.A. Arnone III / Soil Biology and Biochemistry 31 (1999) 517±523
Fig. 3. Aboveground biomass production of plant communities in
1997 as a function of: (a) earthworm density and rainfall
(mean2S.E.) and (b) earthworm activity, in ®eld plots with manipu-
lated earthworm density receiving no additional rain and additional
rain.
signi®cantly between earthworm density treatments
(Fig. 1b).
In a parallel greenhouse study using plant commu-
nities taken from our ®eld site as intact soil-vegetation
monoliths, addition of earthworms resulted in a 15%
increase in aboveground biomass production.
However, this relatively marginal stimulation of plant
biomass production was associated with a 2-fold
greater earthworm density, a 4-fold greater earthworm
biomass and a 10-fold greater earthworm activity (sur-
face cast production; Arnone et al., unpublished data)
than that present in monoliths receiving no additional
worms. These massive earthworm treatments are simi-
lar in magnitude to those imposed in studies reporting
also 10±20%-increases in plant biomass production
after earthworms had been introduced to grasslands
where earthworm abundance was originally very low
(e.g. Stockdill, 1982; Hoogerkamp et al., 1983). Taken
together, these observations suggest: (1) that perennial
plant species common to native grasslands are less re-
sponsive or respond more slowly (e.g. Watkin and
Wheeler, 1966; Chapin, 1980) to earthworm-induced
521
enhancement of nutrient availability than agronomic
plant species, (2) that the signi®cant di€erences in
earthworm activity we created in our ®eld plots by
varying the size of earthworm communities were
apparently insucient to a€ect aboveground plant
growth and (3) that the longer-term e€ects earthworms
have on soil physical properties may be as important,
or even more important, than their shorter-term e€ects
on plant nutrient availability. Finally, there is the
possibility that (4) neither nutrient availability nor soil
structure were constraints to plant growth and were
therefore not alleviated by earthworm activities.
Enhanced aboveground biomass production of plant
communities in plots receiving additional rain was
most likely due to increases in nutrient mineralization
and availability as a result of increased soil moisture
(e.g. Marschner, 1995). These apparent increases in
nutrient availability seem to have been much larger
than the likely increases resulting from enhanced earth-
worm activity in plots with larger earthworm commu-
nities (e.g. Blair et al., 1997).
The lack of a stimulatory e€ect of increased soil
moisture on earthworm activity may be explained in a
couple of ways. First, although the additional rain pro-
vided probably raised soil moisture enough for worms
to remain active, temperatures in the topsoil may have
been prohibitively high (Fig. 2b). Second, earthworm
activity was not a€ected by increased soil moisture
during obligatory worm diapause periods. The lack of
a soil moisture e€ect on earthworm activity further
suggests that the increases in worm activity we
observed in ®eld plots maintained at elevated atmos-
pheric CO2 in another study (Zaller and Arnone,
1997), as well as the increase seen in the greenhouse
study (Arnone et al., unpublished data), was probably
due more to CO2-induced increases in above- and
below-ground detritus (or litter) supply (Arnone,
unpublished data) to worms than to CO2-induced
increases in soil moisture. Our current results showing
a signi®cant increase in the biomass production of
Carex spp. in plots receiving additional rain also indi-
cate that increased soil water availability induced by
elevated CO2 in the ®eld plots (Niklaus and KoÈrner,
1998) may have contributed signi®cantly to the large
increase in Carex aboveground biomass production in
these plots (Leadley et al., 1998).
Thus, our results suggest that moderate increases in
earthworm activity created by manipulating the size of
earthworm communities may not a€ect the plant bio-
mass production in native calcareous grasslands, at
least in the short-term (i.e. two vegetation periods).
Therefore, the apparent e€ects of earthworm activity
on the structure of similar calcareous grassland com-
munities at the ®eld site (Zaller and Arnone, 1998)
seem to be the result of long-term interactions between
plants and earthworms.
522 J.G. Zaller, J.A. Arnone III / Soil Biology and Biochemistry 31 (1999) 517±523
Fig. 4. Aboveground biomass production of graminoids, Carex spp., nonleguminous forbs and legumes in ®eld plots with manipulated earth-
Acknowledgements
We are grateful to Gabrielle Schaer for sorting plant
material. We especially thank Christian KoÈrner for his
comments on an earlier version of the manuscript. The
research was supported by a grant from the Swiss
National Science Foundation to J.A. J.A. is also very
grateful to the Treubel-Fonds of Basel, Switzerland for
its support.
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