Plant and Soil 208: 1–8, 1999.

© 1999 Kluwer Academic Publishers. Printed in the Netherlands.

1

Earthworm responses to plant species’ loss and elevated CO2 in calcareous

grassland

J.G. Zaller1,2,∗ and J.A. Arnone III1,3

1 Botanisches Institut, Universität Basel, Schönbeinstrasse 6, CH-4056 Basel, Switzerland. Present addresses:
2 Ecology Center, Utah State University, 5205 Old Main Hill, Logan, UT 84322-5205, USA; 3 Desert Research
Institute, Biological Sciences Center, 7010 Dandini Blvd., Reno, NV 89512, USA

Received 21 January 1998. Accepted in revised form 29 October 1998

Key words: carbon dioxide enrichment, earthworm electro-sampling, earthworm populations, Lumbricidae, plant
species diversity, plant-animal interactions, soil ecology

Abstract
The objectives of this study were: (1) to quantify the effects of plant species’ loss from designed calcareous
grassland communities at a field site in northwestern Switzerland on the size and composition of earthworm
communities, and (2) to evaluate how exposure of plant communities to elevated atmospheric CO2 might alter the
effects of plant species’ loss on earthworm communities. We non-destructively censused earthworm communities
in each of 24 1.2 m2 experimental plots in autumn 1996 when soils were wet and earthworms were active. Each
plot contained an experimental plant community with 31, 12 or 5 native plant species (eight plots each). Half of
the plots in each species treatment were exposed to ambient CO2 concentrations (350 µL CO2 L−1 ) and half to
elevated CO2 (600 µL CO2 L−1 ) using screen-aided CO2 control. The study was conducted in the fourth year
after community establishment and the third year of CO2 treatment as part of a long-term study on the interactive
effects of plant species’ loss and elevated CO2 on grassland communities. The size (density and biomass) of
earthworm communities declined linearly when the number of plant species in the community was reduced from
31 to 5 species (e.g. 32 ± 1 g m−2 to 23 ± 2 g m−2 ) due mainly to a decline in the endogeic worm species
Allolobophora rosea which was the most abundant of nine earthworm species observed (nearly half of all worms
in each plot). However, no changes in the relative contribution of individual species or the three main earthworm
ecological groups (anecics, endogeics, epigeics) to the entire earthworm community were observed with declining
number of plant species. The responses of earthworm communities to plant species’ loss appear to reflect changes
in community fine root biomass in the topsoil (e.g. declining worm biomass with declining fine root biomass)
observed in parallel studies conducted at this site. Further the results of this study demonstrate that a loss of plant
species’ from these calcareous grassland communities may also alter the age structure of earthworm communities,
but not significantly influence their diversity or composition. Our data also indicate that rising atmospheric CO2
may not greatly impact the size and composition of worm communities or alter the effects of plant species’ loss on
earthworm communities. Therefore, the disappearance of plant species from these native grasslands, as a result of
ever increasing human activities, may be expected to lead to reductions in the size of earthworm communities and
the ecosystem services they provide.

Introduction ing to the rise of atmospheric CO2 levels, are expected

Human activities leading to loss of plant species from
natural terrestrial ecosystems, and activities contribut-
∗ FAX No: +1435 797 3872. E-mail: jgz@cc.usu.edu
to alter the functioning of these systems globally (e.g.
Chapin et al. 1997, Hooper and Vitousek, 1997;
Körner, 1995; Peters and Lovejoy, 1992; Schulze and
Mooney, 1993; Wedin and Tilman, 1990). Soil het-
erotrophs are key members of these ecosystems who,
2
as decomposers and nutrient cyclers, not only depend
on plants as primary producers but who also feed back
to affect plants (e.g. Coleman and Crossley, 1996).
Thus, any changes in the community of primary pro-
ducers, resulting from either species’ loss or induced
by elevated CO2 , must affect soil heterotrophs, and
thus the functioning — and perhaps even the stability
(e.g. McNaughton, 1977; Tilman et al., 1996) and
resilience (e.g. Tilman, 1996) — of the ecosystem.
However, experimental data needed to test these asser-
tions are scant, and even less is known about how plant
species’ loss and rising atmospheric CO2 will interact.
In calcareous grasslands of central Europe, earth-
worms are among the most important soil heterotrophs
(70 – 80% of soil zoomass). At our field site, they
are responsible for an annual turnover of about 16 t
soil (dry mass) ha−1 and 70 kg N ha−1 under cur-
rent ambient CO2 concentrations, and up to 35%
more in grassland plots maintained under elevated at-
mospheric CO2 (Zaller and Arnone 1997). Increased
activity under elevated CO2 appears mainly to be due
to enhanced carbon inputs to the soil (cf. Zaller and
Arnone, 1998a) but also to CO2 -induced increases in
soil moisture (Zaller and Arnone, 1997). It is uncer-
tain, however, whether increased activity was due to
larger earthworm communities (as reported by Yeates
et al. (1997) in managed New Zealand pasture land),
or simply enhanced feeding and burrowing activity.
Loss of plant species from these calcareous grass-
lands may impact earthworm communities in two
important ways: first, via changes in net primary
productivity (NPP; sensu Tilman, 1996); and sec-
ond via the elimination of certain plant species to
which earthworms are particularly attracted (because
of the palatability of their litter or because of the plant
species’ effect on stand microclimate; see Zaller and
Arnone 1998b). In the designed plant communities
established at our site in which the number of native
plant species present was reduced in three steps from
the present level of 31 to 12 to 5 (Leadley and Körner,
1996), plant species’ loss did not affect net ecosys-
tem carbon uptake (NEC; Stocker et al., 1998) or net
aboveground biomass production (P. Leadley, personal
communication). This suggests that plant species’ loss
may not affect NPP significantly. Fine root biomass
in the topsoil, on the other hand, declined signifi-
cantly with reductions in plant species (P. Leadley,
personal communication) suggesting that carbon sup-
ply to soils may decline with elimination of plant
species from communities. Consequently, the size or
activity of earthworm communities may also be re-
duced by plant species’ loss. Further, the functional
composition of earthworm communities, viewed at
either the species or ecological-group level (anecic,
endogeic and epigeic species), may even be altered.
In field plots where species’ number was manip-
ulated, elevated CO2 caused several important effects
relevant to our study of earthworm communities. First,
soil moisture was higher in all plots maintained at
high CO2 (P. Niklaus, unpublished data; Zaller and
Arnone, 1997) theoretically improving the environ-
ment for earthworms. Second, NEC (Stocker et al.,
1998) was markedly greater in communities main-
tained under elevated CO2 , than in plots maintained at
ambient CO2 , but plant biomass only slightly higher
(P. Leadley, pers. comm.) indicating an increase in
carbon inputs to the ecosystem.
The objectives of our study were to quantify
the effects of a loss of plant species and elevated
atmospheric CO2 on (1) the overall size (num-
bers/density and biomass) of earthworm communities,
(2) individual earthworm species’ responses (density
and biomass), (3) the representation of species and
the three main earthworm ecological groups, (4) the
earthworm species’ diversity, and (5) the age structure
of earthworm communities.
Materials and methods
Study site and experimental design
This study was conducted as part of an ongoing, long-
term study designed to evaluate the effects of plant
species composition and elevated CO2 on the ecology
of calcareous grasslands (e.g. Körner, 1995; Leadley
and Körner, 1996). The study site is located near
the village of Nenzlingen south of Basel on a 20◦
southwest-facing slope in NW-Switzerland (520 m,
47◦ 27’ N, 7◦ 34’ E). Average annual precipitation is
about 900 mm, average annual air temperature be-
tween 8.5–9.0 ◦ C (Ogermann et al., 1994). Soils are
classified as a Rendzina (pH is about 6.5, bulk density
of the top soil 1.1 g/cm3 ), with a well developed, rock-
free, loamy topsoil and a rapid transition at 10–15 cm
depth to the underlying scree material (Ogermann et
al., 1994). Up to 1993 this calcareous grassland had
been used as an extensively managed pasture, mainly
for cattle.
The overall experimental setup and design is de-
scribed in detail in Leadley and Körner (1996). Briefly,
the topsoil from 24 hexagonal plots (1.2 m2 ) was

Table 1. Number of plant species per functional group
used to construct grassland communities. Half of the
communities were exposed to ambient atmospheric
CO2 concentrations (350 µL CO2 L−1 ) and half to
elevated CO2 (600 µ CO2 L−1 ); n = 4. Abstracted
from Leadley and Körner (1996)
Functional groups Number of plant species
High Medium Low
Graminoids 8 2 1 V DC) was applied over the entire length of the rods
Non-legume forbs 17 7 2
Legumes 6 3 2
Total number of spp. 31 12 5
removed in the spring of 1993, homogenized and
returned to the plots. Most plant species used to estab-
lish communities were propagated from seed collected
from the site. Some species were vegetatively propa-
gated using local material. Communities were created
by planting individuals from either 31, 12 or 5 species
on a hexagonal grid with a distance of 3.8 cm between
individuals (590 individuals m−2 ). The representation
of each of the three functional groups — graminoids
(55% of all individuals planted), non-legume forbs
(30%) and legumes (15%) — was held constant across
all plots, with only the number of species in each
functional group varying (Table 1). Since March 1994,
four plots of each community type have been main-
tained round the clock (except from December to early
March during winter months) to elevated CO2 concen-
trations (600 µL CO2 L−1 ) and half to ambient CO2
concentrations (350 µL CO2 L−1 ) using screen-aided
CO2 control (SACC: 50-cm-tall transparent open-top,
open-bottom chambers; Leadley et al., 1997). Plots
were arranged in a randomized complete block design
with four blocks.
Earthworm sampling
Earthworms in each plot were censused in October
1996 when soils were wet and earthworms were active
(i.e. based on surface cast production) using the so-
called ‘Octet’ electro-sampling method (Thielemann,
1986). We chose to use this method for two main
reasons. First, we needed to insure that neither earth-
worms nor plants in the community would be damaged
by repeated sampling required to census earthworm
communities. Second, this non-destructive method has
been shown to provide comparable estimates of earth-
worm community size and composition to other more
3
conventional sampling methods (hand-sorting and for-
maline infiltration methods), as long as earthworms
are sampled at times when they are active and when
soil moisture is sufficient (Vetter, 1996).
Electro-sampling involved pushing eight steel rods
(65 cm long × 6 mm dia.) into the ground in the centre
of each plot along a circumference of a 60 cm diameter
circle at equal distances. Electrical voltage (300–500
in 4-second pulses supplied pairwise to opposing rods
in the ‘Octet’. Each successive electrical voltage was
applied to the next pair of opposing rods. The portable
transformer (DEKA 4000, Deka Gerätebau, Germany)
providing electrical pulses was powered by a 12 V
car battery. All earthworms driven to the soil sur-
face within 40 min were collected and stored in cool
water. Earthworms were sorted by species (Bouché,
1972) and age class (adult: well developed clitellum;
subadult: developed puberculum, but no clitellum; ju-
venile: puberculum and clitellum absent), assigned
to one of three ecological groups (epigeics, anecics,
endogeics; Bouché, 1977), and were counted and
weighed. Epigeic earthworm species live in or near the
surface litter feeding primarily on coarse particulate
organic matter. Endogeic species live within the soil
profile and predominantly feed on soil and associated
organic matter, whereas anecic species live in vertical
burrow systems and are believed to feed primarily on
surface litter which they pull into their burrows (e.g.
Bouché, 1977; Edwards and Bohlen, 1996).
Statistical analysis
The General Linear Model-ANOVA procedure in the
SYSTAT statistical package (SYSTAT, v. 5.2.1, Sys-
tat Inc., Evanston, IL, USA; Wilkinson et al., 1992)
was used to test the effects of Plant Species Num-
ber (df = 2), CO2 (df = 1), Block (df = 3) and
Plant Species Number × CO2 (df = 2) on: commu-
nity earthworm density and biomass, the density and
biomass of individual earthworm species common to
all plots (six species), the relative contribution of in-
dividual species and ecological groups to community
density and biomass, earthworm community diver-
sity (Shannon index, see below), and the contribution
of earthworm age classes to earthworm community
density and biomass. Relative contribution data were
arcsine-transformed prior to statistical analyses (Sokal
and Rohlf, 1981). The Shannon diversity index (cf.
Zar, 1996) was calculated for each plot using e as
logarithmic base to assess the effects of treatments on
4
earthworm community diversity. In the interpretation
of our results we considered P < 0.05 as significant
and 0.05 < P ≤ 0.10 as marginally significant. Treat-
ment means given in the text and figures are presented
with their standard errors (±SE).
Results
Total density and biomass of earthworm communities
declined linearly with declining plant species number
(density: from 106 ± 11 to 73 ± 20 worms m−2 ;
biomass: from 31.8 ± 1.3 to 22.5 ± 2.3 g fresh
mass m−2 ), however the decline in density was only
marginally significant (P = 0.108; Figure 1). Elevated
CO2 had no apparent effects on either earthworm
community density or biomass (Figure 1).
We found nine species of earthworms across all
treatments, six of which occurred in all plots. Anecic
species (Lumbricus terrestris L., Nicodrilus longus
Ude, Nicodrilus nocturnus Ude) accounted for 35%
of total earthworm community biomass. Endogeic
species (Octolasion cyaneum Sav., Allolobophora
rosea Sav., A. chlorotica Sav., Nicodrilus caligi-
nosus Sav.) represented the largest ecological group
with 62% of total earthworm biomass, while epigeic
species (Lumbricus castaneus Sav., Dendrobaena
mammalis Ger.) represented the smallest fraction with
3% of total biomass. Nicodrilus nocturnus, and Lum-
bricus terrestris were present in only two of the 24
plots, and Dendrobaena mammalis in only one. Al-
lolobophora rosea was the most abundant earthworm
species in all plots (49% of all worms averaged across
treatments), followed by Octolasion cyaneum (19%),
Allolobophora chlorotica (11%), Nicodrilus longus
(10%), Lumbricus castaneus (6%) and Nicodrilus
caliginosus (5%) (Figure 2).
Overall, the density and biomass of individual
earthworm species and earthworm ecological groups
was affected very little by either plant species’ loss or
elevated CO2 (Figure 2). Consequently, the contribu-
tion of individual earthworm species, and earthworm
ecological groups, remained largely unaffected by
these treatments (P = 0.40, data not shown). Plant
species’ loss appeared to only impact the endogeic
species Allolobophora rosea and Nicodrilus caligi-
nosus as indicated by a decline in their populations
(density and biomass; Figure 2). The decline in these
two species, but especially in A. rosea, contributed
substantially to the overall decline in earthworm com-
munity biomass described above. The only significant
effect of elevated CO2 appeared to be a reduction in A.
rosea biomass. Neither the loss of plant species from
communities nor elevated CO2 affected the diversity
of earthworm communities(Table 2).
Table 2. Shannon diversity index of the earthworm com-
munity as a function of the number of plant species in a
community and atmospheric CO2 concentrations (ambient
CO2 ; 350 µL CO2 L−1 ; elevated CO2 ; 600 µL CO2 L−1 ;
mean ± SE, n = 4)
CO2 Number of plant species in communities
31 12 5
Ambient 1.27 ± 0.08 1.28 ± 0.08 1.50 ± 0.15
Elevated 1.25 ± 0.05 1.28 ± 0.10 1.31 ± 0.12
The age structure of all earthworm commu-
nities was hour-glass-shaped, where 80% of all
worms viewed across treatments were juveniles, 2%
subadults, and 18% adults (Figure 3). Reductions in
the number of plant species affected only the contribu-
tion of subadult earthworms to the entire earthworm
community, resulting in a linear decrease from 5%
in communities with 31 plant species to 1% in com-
munities with 5 plant species. Elevated CO2 did not
alter this pattern, nor did it affect the contribution of
adult worms to the community. However, the effects
of declining number of plant species on the contribu-
tion of juvenile earthworms to the worm community
were different in communities maintained at ambient
and elevated CO2 (Figure 3).
Discussion
The reduction in earthworm community biomass and
density by about 30% with reduction in the num-
ber of plant species from 31 to 5 in these calcareous
grassland communities four years after plant commu-
nities were established paralleled reductions in root
biomass observed in the topsoil of these communi-
ties with increasing species’ loss (P. Leadley, pers.
comm.). This suggests that plant detritus made avail-
able to earthworms via the fine roots (e.g. rhizode-
position or support of rhizosphere microorganisms
consumed by earthworms) regulates earthworm com-
munity size (density and biomass). The apparent linear
decrease in worm community size with reductions
in the number of plant species (Figure 1) suggests

Figure 1. Density (a) and biomass (b) of earthworm communities as a function of the number of plant species present in a community (31, 12,
and 5 species) and the atmospheric CO2 concentration (ambient CO2 : 350 µL CO2 L−1 , and elevated CO2 : 600 µL CO2 L−1 ). Mean ± SE,
n = 4. P values derive from ANOVAs with Plant Species Number (Pl), CO2 concentration (CO2 ), and Block as factors.
that each plant species plays an equal role (perhaps
via its contribution to plant community fine root bio-
mass) in sustaining earthworm communities as long as
the contribution of plant functional types to the plant
community is held constant.
Significantly increased ecosystem carbon uptake
by all communities maintained under elevated at-
mospheric CO2 at our site (Stocker et al., 1998; but
also reported in other studies: Hungate et al., 1997;
Tate and Ross, 1997), as well as increased soil mois-
ture under elevated CO2 (P. Niklaus, unpublished data;
Zaller and Arnone, 1997), had no apparent effect on
the overall size of earthworm communities, even af-
ter three years of CO2 treatment. This is surprising
because we have observed dramatic increases in earth-
worm activity (surface casting) in calcareous grassland
ecosystems maintained under elevated CO2 (Arnone et
al., 1998; Zaller and Arnone, 1997), including in the
plots containing 31 plant species used in the present
study (G. Hofer, unpublished data). Taken together
these data suggest that elevated CO2 only stimulates
the activity of earthworms and does not necessarily
alter the size of their populations (see also Yeates et
al., 1997). Additionally, the lack of any significant
differences in topsoil fine root biomass between com-
munities maintained at ambient and elevated CO2 (P.
Leadley, pers. comm.), further supports the notion that
some property associated with fine roots is important
for maintaining earthworm communities.
5
Although plant species’ loss and elevated CO2 had
relatively little effect on the composition of earthworm
communities at either the ecological-group or species
level, their suppressive effects on endogeic species,
particularly A. rosea, may have wide ecological ramifi-
cations. This is because earthworm communities in the
plots we studied are made up mostly (62%) of species
from this ecological group known to be important
in the establishment and maintenance of soil struc-
ture (e.g. Bolton and Phillipson, 1976; Scheu, 1987;
Springett, 1983). The absence of any treatment effects
on earthworm community diversity suggests that re-
ductions in the diversity of communities of primary
producers may not affect the diversity of associated
heterotrophs and that rising CO2 may not alter this.
The overall age structure of earthworm communi-
ties censused in autumn 1996, with almost four times
more juveniles than subadults and adults together, was
typical of that exhibited by soil-inhabiting inverte-
brates (e.g. Edwards and Bohlen, 1996; Gerard, 1967;
Lee, 1985). In general, the fact that both plant species’
loss and elevated CO2 affected the age structure of
earthworm communities suggests that these two im-
portant global changes may also affect earthworm
population and community dynamics.
Thus, the results of this study demonstrate that a
loss of plant species’ from these calcareous grassland
communities may: (1) result in reductions in the size
of earthworm communities, due to reductions in cer-
tain endogeic species; and (2) alter the age structure of
6

Figure 2. Density and biomass of earthworm species in grassland communities with different plant species numbers (31, 12, and 5 species)
and atmospheric CO2 concentration (ambient CO2 : 350 µL CO2 L−1 , open symbols; elevated CO2 : 600 µL CO2 L−1 , filled symbols). Mean
± SE, n = 4. Earthworm species arranged by ecological group. Note the use of different scales on Y-axes. P values derive from ANOVAs with
Plant Species Numbers (Pl), CO2 concentration (CO2 ), and Block as factors. Treatment effects at the P < 0.10 are shown in bold-faced type.

Figure 3. Age structure of earthworm communities (e.g. contri-
bution of juveniles, subadults, and adults to the total density of
earthworm communities) in grassland communities as a function of
plant species number (31, 12, and 5 species) and atmospheric CO2
concentration (ambient CO2 : 350 µL CO2 L−1 , elevated CO2 : 600
µL CO2 L−1 ). Mean ± SE, n = 4.
7
earthworm communities; however (3) not significantly
influence the diversity or composition of earthworm
communities. Our data also indicate that rising at-
mospheric CO2 may not greatly impact the size and
composition of worm communities or alter the ef-
fects of plant species’ loss on earthworm communities.
Therefore, the disappearance of plant species from
these native grasslands, as a result of ever increasing
human activities, may be expected to lead to reduc-
tions in the size of earthworm communities and the
ecosystem services they provide.
Acknowledgements
We are grateful to Paul Leadley and Pascal Niklaus
for maintaining the field CO2 enrichment facility, and
to Christian Körner and P. N. for reviewing an earlier
version of the manuscript. Beate Kittl and Monika
Keller helped with earthworm sampling. This work
was funded by grants from the Swiss National Sci-
ence Foundation to J. A. A. III (NF 3100-042401.94/1)
and Ch. K., J. A. A. III and Bernhard Schmid (NF-
SPPU 5001-035214). J. A. A. III greatly appreciates
the generous support of the Treubel-Fonds of Basel,
Switzerland.
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Section editor: E Garnier