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www.elsevier.com/locate/agee
Agriculture, Ecosystems and Environment 109 (2005) 323–334
Effects of fertiliser and grazing on the arthropod
communities of a native grassland in
south-eastern Australia
Ian Oliver a,*, Denys Garden b, Penelope J. Greenslade c,Bronwyn Haller a, Denis Rodgers d, Owen Seeman e, Bill Johnston f
a New South Wales Department of Infrastructure, Planning and Natural Resources, PO Box U245,
University of New England, Armidale, NSW 2351, Australiab New South Wales Department of Primary Industries, GPO Box 1600, Canberra, ACT 2601, Australia
c Division of Botany and Zoology, Australian National University, Canberra, ACT 0200, Australiad 19 Amaroo Avenue, Ferny Hills, Qld 4055, Australia
e PO Box 3300, The Queensland Museum, South Brisbane, Qld 4101, Australiaf New South Wales Department of Infrastructure, Planning and Natural Resources,
PO Box 365, Queanbeyan, NSW 2620, Australia
Received 13 January 2004; received in revised form 18 February 2005; accepted 21 February 2005
Abstract
An experiment commenced in 1998 to test the effects of superphosphate fertiliser application and grazing on production
and botanical composition of a native grassland in south-eastern Australia. Superphosphate application resulted in an
increase in sheep production but a decline in native perennial grasses and an increase in exotic annual grasses. The study
reported here aimed to determine if arthropod assemblages showed changes in community composition on the same
experimental plots. The experiment was conducted in grassland dominated by the native perennial wallaby grass,
Austrodanthonia duttoniana, and consisted of six replicated treatments that were designed to improve grassland and
domestic livestock productivity. Treatments consisted of a control (no fertiliser), three levels of annually applied super-
phosphate (62.5, 125, and 250 kg ha�1), and two treatments aimed to raise soil pH (superphosphate plus lime, and sewage
ash). Soil properties were measured annually and sheep stocking rates were increased over the duration of the experiment
according to increases in available forage. Soil and ground-active arthropod populations were sampled from all plots in
spring 2001. Fertiliser application and grazing increased the relative abundance of introduced Acari and Collembola, and
changed the community composition of Formicidae and Coleoptera. Lime and sewage treatments had variable effects on
taxa. Improving the productivity of native grassland with superphosphate led to a decline in plant and arthropod biodiversity
through reduced abundance and/or local extinction of native species and increased dominance of introduced species. These
* Corresponding author. Tel.: +612 6773 5271; fax: +612 6773 5288.
E-mail address: [email protected] (I. Oliver).
0167-8809/$ – see front matter. Crown Copyright # 2005 Published by Elsevier B.V. All rights reserved.
doi:10.1016/j.agee.2005.02.022
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334324
findings support the need to protect and restore a representative network of native grassland ecosystems within the
agricultural zone of south-eastern Australia.
Crown Copyright # 2005 Published by Elsevier B.V. All rights reserved.
Keywords: Native grassland; Fertiliser; Lime; Sewage; Grazing; Arthropod biodiversity; Acari; Collembola; Coleoptera; Formicidae; Exotic
species
1. Introduction
Within Australia’s Murray-Darling Basin the major
land use is domestic livestock grazing. Because
Australian soils are relatively low in phosphorus
(P), sulphur (S) and nitrogen (N) (King and
Hutchinson, 1983), native grassland may be improved
through the addition of P and S as superphosphate, and
N fixed by sown legumes (King et al., 1985). To raise
the pH of soils and increase productivity, liming has
also become more common in recent years.
Native grasslands and their soils contain large
numbers of arthropods, the biodiversity of which is
vitally important to production systems (Srivastava
et al., 1996; Vietmeyer, 1996; Altieri, 1999; Paoletti,
1999; CSIRO, 2001). The impacts of fertiliser
application and associated increases in stocking
intensity on arthropod biodiversity can vary by
location and taxa (Zyromska-Rudzka, 1979; King
and Hutchinson, 1980; King et al., 1985; Berger et al.,
1986; Carlyle and Than, 1987; Vickery et al., 2001). In
Australia, King and Hutchinson (1980) found an
increase in mite and collembolan populations follow-
ing superphosphate application. Zyromska-Rudzka
(1979) found that scutacarid mites decreased in
abundance after NPK fertilisation of a Polish meadow,
while oribatid mites increased, whereas Olechowicz
(1977) found application of NPK fertiliser to result in
an increase of dipterans. Carabid beetles and spiders
decreased in abundance and species richness follow-
ing fertiliser application (Luff and Rushton, 1989;
Rushton et al., 1989). King et al. (1985) found that the
relative abundance of native and introduced arthropod
species changes, with native species of Collembola
dominating native grassland, but introduced or
cosmopolitan species dominating in fertilised pastures
sown to exotic species.
Similarly, arthropods may respond to grazing
induced changes in habitats in a variety of ways.
For example, disturbance-tolerant beetles in the
families Scarabaeidae and Curculionidae (Kain,
1979; Bromham et al., 1999), and ants in the genera
Iridomyrmex and Rhytidoponera (Hutchinson and
King, 1980a; Lobry de Bruyn, 1993; Bromham et al.,
1999; Read and Andersen, 2000) were found to
increase in abundance following grazing, whereas,
other less tolerant groups declined (Morris, 1968;
King and Hutchinson, 1976, 1980, 1983; King et al.,
1976; Hutchinson and King, 1980a,b; Bromham et al.,
1999; Seymour and Dean, 1999).
This study of arthropod populations adds a further
dimension to an experiment that commenced in 1998,
in which different rates of superphosphate fertiliser
and other soil amendments were applied to a native
grassland in south-eastern Australia, with the aim of
improving grassland and domestic livestock produc-
tivity (Garden et al., 2003). The study found that
superphosphate application and increased grazing
resulted in a decline in native perennial grasses and an
increase in exotic annual grasses. Our study used the
same experimental plots and asked the question, do
arthropod assemblages within a native grassland show
changes in community composition in response to
increased fertiliser and grazing? The term grassland is
used rather than pasture to indicate that the experi-
mental area had not been sown to exotic species or
disturbed by cultivation or the use of herbicides.
2. Methods
The study site was located 22 km north of Yass on
the southern tablelands of New South Wales, Australia
(348400S 1488540E, altitude 670 m). Soils were
derived from Silurian tuffs, and varied from shallow
(<20 cm) Leptic Rudosol soils through bleached
mottled Chromosol and yellow Sodosol soil to deep
(>1.5 m) Hydrosol soils in low-slope areas. Soils were
low in P and S, and acidic in the surface layers
(pHCaCl2 4.1–4.9). Annual rainfall was 668 mm
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334 325
evenly distributed throughout the year. Before treat-
ments the site was dominated by native C3 species,
mainly Austrodanthonia duttoniana (Cashmore) H.P.
Linder. Introduced annual species (Vulpia spp. and
Bromus spp.) were present at low densities, with some
Trifolium subterraneum L. and other annual clovers
(total species 64; native species 45% by number, 77%
by mass, Garden et al., 2003). No trees occurred in
study plots. The site had previously been grazed by
sheep at low-medium rates (3–5 sheep/ha) with
limited fertiliser (mainly P) application.
Experimental plots ranged from 0.5 to 0.83 ha with
two replicates of each of six treatments. Treatments
were randomly allocated to plots within blocks either
side of a central access lane, with replicate 1 (plots
1–6) generally being higher in the landscape and
having shallower soils. Treatments consisted of: a
control with no improvement; three levels of super-
phosphate (8.8% P, 11% S), i.e. low (62.5), moderate
(125) and high (250 kg ha�1); lime (250 kg ha�1
superphosphate + 2.5 t ha�1 lime); and sewage (5 t
ha�1 sewage ash). Superphosphate treatments were
applied in March 1998, 1999, May 2000, April 2001
and March 2002. Lime and sewage treatments were
applied in March 1998 only. Molybdenum was applied
as molybdenised superphosphate (0.02% Mo) to all
treatments (except sewage) in 1998 and 2002 at
25 g ha�1. Stocking rate was varied according to
available forage within treatments (Table 1), with the
aim of maintaining similar body weights across
treatments, and was adjusted by adding wethers from
a mob of similar animals grazed on similar unfertilised
pasture.
Table 1
Stocking rate (wethers/ha) as determined by available forage at the
experimental site in NSW
Year Month Control Low Moderate High Lime Sewage
1998 May 6 6 8 8 8 8
October 6 7.2 9.6 10 10 8
December 6 7.2 9.6 12 12 10
1999 October 7.2 8.4 11.2 14 14 12
2000 March 7.2 8.4 11.2 14 14 14
May 8.4 9.6 12.8 16 16 16
2001 May 8.4 10.8 14.4 16 20 16
Control: no fertiliser, low: 62.5, moderate: 125, high: 250 kg ha�1
superphosphate, lime: 250 kg ha�1 superphosphate + 2.5 t ha�1
lime, sewage: 5 t ha�1 sewage ash.
Soils were sampled before the experiment and
in autumn each year before fertiliser was applied.
From each plot 20 soil samples were collected 10 cm
deep along two separate 40 m transects and bulked
for analysis. They were dried, ground to pass a
2 mm sieve, and analysed by a commercial soil
testing laboratory for electrical conductivity, pHCaCl2 ,
PColwell, exchangeable cations, SKCl40 and organic
carbon, the latter two in autumn 2002 only.
Herbage mass (kg dry matter ha�1), ground cover
and botanical composition (percent of herbage mass)
were assessed every 6–7 weeks within 30 quadrats
(0.5 m � 0.5 m) along the same transects for soil
sampling using the BOTANAL technique (Andrew
and Lodge, 2003; Garden et al., 2003) and used to
determine the proportion of herbage mass contributed
by exotic species.
Sheep were removed from the plots in spring 2001
during arthropod sampling. Arthropods active on the
ground surface (epigaeic) were sampled using ten
250 ml pitfall traps (100 mm high, 67 mm internal
diameter), buried with the lip of the container flush
with the soil surface. Each contained 125 ml mono-
ethylene glycol to kill and preserve the trapped
arthropods. On November 6, 2001, two rows of pitfall
traps were installed in each plot 5 m either side of a
central 50 m transect. Traps were located at 5, 15, 25,
35, and 45 m along this transect and were active for 7
days after which they were removed, capped and
transported to the laboratory where the monoethylene
glycol was decanted off and replaced with 70%
ethanol.
Arthropods active within the soil (hypogaeic) were
recovered from soil cores taken within 50 cm of each
pitfall trap. Cores of 7.5 cm diameter, 5 cm deep were
taken through the grass, wrapped in aluminium foil
and transported to the laboratory for arthropod
extraction into 100% ethanol over 7 days using
Tullgren funnels (15 W globe 20 cm from the inverted
soil core). Five soil cores from each replicate were
taken on two occasions, the first when pitfall traps
were installed, the second when they were removed. A
random set of nine soil core samples was selected from
each replicate. Pitfall trap and soil core samples were
pooled per plot prior to identification and analysis.
Arthropods were identified to orders, morphospe-
cies (Oliver and Beattie, 1996a,b) or species, and
categorised into native or exotic taxa where possible.
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334326
Table 2
Abundance and richness of arthropods sampled by pitfall traps at the
experimental site in NSW
Abundance Richness
Formicidae 619 (0.11) 13 (0.11)
Voucher specimens were deposited with the Austra-
lian Museum, Sydney (Formicidae, Coleoptera and
Hemiptera), the Australian National Insect Collection,
Canberra (Acari) or the South Australian Museum,
Adelaide (Collembola).
Coleoptera 281 (0.05) 43 (0.38)Collembola 1688 (0.30) 23 (0.20)
Acari 1744 (0.31) 28 (0.25)
Hemiptera 169 (0.03) 7 (0.06)a
Diptera 732 (0.13)b
Others 394 (0.07) –
Total 5627 114
a Hemiptera data were not analysed further due to low numbers of
species.b Diptera were not identified to species nor analysed further.
3. Analyses
Differences in total abundance and relative
abundance of exotic arthropods among treatments
were explored graphically and tested using one-way
anova. Changes in the soil and vegetation parameters
due to treatments were explored graphically and tested
using two-way anova (Gmav5, Underwood, 1995).
The ability of soil and vegetation parameters to
explain the variation in arthropod assemblages in
spring 2001 was tested using the direct gradient
analysis technique canonical correspondence analysis
(CCA) (CANOCO for Windows, ter Braak and
Smilauer, 2002). Correlated variables were first
identified by significant simple correlation coefficients
using data from final sampling periods and by
inspection of variance inflation factors within initial
CCAs (ter Braak and Prentice, 1988). Final CCAs
excluded correlated variables and were performed on
raw and transformed (x0.25) data. All default settings
were used for CCA other than requesting 9999 Monte
Carlo permutations to test for statistically significant
explanatory variables. Variables were tested indivi-
dually, and for their ability to explain significant
remaining variation in minimum models built by
forward selection. Tri-plots were produced using
CanoDraw (ter Braak and Smilauer, 2002).
4. Results
The pitfall traps sampled 5627 specimens from 114
arthropod species during the 7-day period in spring
2001 (Table 2), and the soil cores sampled 21,405
sarcoptiform mites from 15 species, and 5997
Collembola from 18 species. Collembola and Acari
were more abundant at higher superphosphate applica-
tion, Formicidae and Coleoptera in the control,
abundance under lime and sewage showing variable
trends among groups (Fig. 1). Average relative
abundance of Collembola and Acari of exotic origin
increased with increasing fertiliser application and
grazing (Fig. 2), statistically so for the hypogaeic Acari,
where the exotic Tyrophagus similis Volgin represented
96.7% of the sarcoptiform mites recovered.
Two-way anova revealed significant interaction
between treatment and time for pH (P < 0.001), and
phosphorus (P < 0.01), demonstrating that soil pH and
P had changed during the experiment and to varying
degrees among treatments (Fig. 3). Significant treat-
ment effects were also obtained for ground cover, exotic
biomass (Fig. 3) and S (one-way anova P < 0.01, S
treatment means were: control 3.5, low 3.9, moderate
5.9, high 7.8, lime 11.3, and sewage 3.4).
Simple correlations between February 2002 soil
data (d.f. = 10, P < 0.05 when r > 0.576) revealed a
number of highly correlated soil variables including:
EC, correlated with Al, Ca, K, Mg, Na, pH and S
(r = �0.810, 0.977, 0.669, 0.625, 0.586, 0.907, and
0.701, respectively); Al, correlated with Ca, K, P and
pH (r = �0.897, �0.653, �0.622, �0.951); and Ca,
correlated with K, pH and S (r = 0.658, 0.969, 0.643).
The three collinear variables EC, Al, and Ca were
therefore omitted from further analysis.
Initial CCAs (with EC, Al and Ca excluded)
revealed high variance inflation scores attributable to
S. Sulphur, which was positively correlated with K,
Na, pH and exotic biomass (r = 0.747, 0.693, 0.590,
and 0.588, respectively), was therefore omitted from
the final CCAs. All variance inflation factors for final
CCAs were less than 20. Final CCAs based on raw
data revealed that, when tested individually, ground
cover and exotic biomass explained significant
(P < 0.05) variation in hypogaeic Acari and hypo-
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334 327
Fig. 1. Average total abundance of arthropods by treatment (with S.E.; Student–Newman–Keuls post-hoc multiple comparison tests).
gaeic Collembola assemblages (Table 3). However,
forward selection of variables found that exotic
biomass did not explain significant remaining varia-
tion in the models containing ground cover alone,
which accounted for 25% and 22% of the variation in
hypogaeic Acari and Collembola assemblages,
respectively. Exotic biomass was the single statisti-
cally significant explanatory variable for the epigaeic
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334328
Fig. 2. Average relative abundance of arthropods of exotic origin by treatment (with S.E.; Student–Newman–Keuls post-hoc multiple
comparison tests).
Acari and accounted for 22% of the variation. In
comparison, forward selection of variables found that
ground cover, pH and organic carbon together
accounted for 61% of the observed variation in the
epigaeic Collembola assemblage. A number of other
variables, including exotic biomass, were individually
significant but were not included in the minimum
model (Table 3). Organic carbon and ground cover
accounted for 45% of the observed variation in the
Formicidae assemblage, and soil P was the single
statistically significant explanatory variable for the
Coleoptera assemblage and accounted for 14% of the
variation. Results from CCAs based on transformed
(x0.25) abundance data were similar.
CCA tri-plots (Fig. 4) revealed separation of
control and low treatment plots from moderate and
high treatment plots, the former characterised by
lower ground cover and lower exotic plant biomass,
and by the native Collembola: Brachystomella
platensis Najt and Massoud (NU1), Proisotoma sp.
(N17), Subisotoma sp. (N9), and Corynephoria sp.
(N22) (Fig. 4a and c); and the native Acari:
Zygoribatula cycloporosa Lee (N3), Scheloribates
brevipodus Lee and Pajak (N19), Lanceoppia tasma-
nica Mahunka, (N1) Anellozetes sp. (N4), Rhagidia
sp. (N10), as well as the exotic species Petrobia latens
(Muller) (E4) (Fig. 4b and d). The higher treatment
plots were characterised by higher ground cover,
higher exotic plant biomass, the exotic Collembola:
Hemisotoma thermophila (Axelson) (E1), Hypogas-
trura vernalis (Carl) (E5), Entomobrya multifasciata
(Tullberg) (E2) and Isotomurus sp. (E3) (Fig. 4a and
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334 329
Table 3
Variables that accounted for significant variation in arthropod assemblages when tested by CCA individually (ind) or as part of a minimum model
(mod)
Variable Acari Collembola Formicidae Coleoptera
Epigaeic Hypogaeic Epigaeic Hypogaeic Epigaeic Epigaeic
Ground cover ns mod, ind mod, ind mod, ind mod, ind ns
Exotic biomass mod, ind ind ind ind ns ns
Organic carbon ns ns mod ns mod, ind ns
pH ns ns mod, ind ns ns ns
Phosphorus ns ns ind ns ns mod, ind
Potassium ns ns ind ns ns ns
Variation explained (%)a 22 25 61 22 45 14
a Variation explained by minimum model calculated as the sum of all canonical eigenvalues divided by sum of all unconstrained eigenvalues
when the model included only those variables contained within the minimum model. Significance was tested by Monte Carlo permutation tests
with 9999 permutations, ns P > 0.05.
Fig. 3. Changes in some soil and vegetation parameters by treatment during the experiment (average of two plots per treatment are plotted). Plots
are identified by their treatments as: Control (&), Low ( ), Moderate ( ), High ( ), Lime ( ), Sewage (^). Two-way anova results for sources of
variation: treatment (tr), time (ti), and treatment x time interaction (txt), are *P < 0.05, **P < 0.01, ***P < 0.001. (a) pH, tr*** ti** txt***, (b)
phosphorus, tr** ti*** txt**, (c) ground cover, tr*, (d) exotic biomass, tr*** ti***.
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334330
Fig. 4. Tri-plots based on canonical correspondence analysis of Collembola and Acari raw abundance data. First (horizontal) and second
(vertical) canonical axes are plotted over the range �1 to +1. Vectors are shown as: dotted, when P > 0.05 for marginal test (variables tested
individually with 9999 permutations); solid when P < 0.05 for marginal test; and bold when P < 0.05 for conditional test (variables added to
minimum model by forward selection). Plots are identified by their treatments as: Control (&), Low ( ), Moderate ( ), High ( ), Lime ( ),
Sewage (^). Taxa are shown as alphanumeric couplets with E = exotic, N = native, NU = probable native taxa, and U = taxa of unknown origin.
All taxa were submitted to CCA but only those recorded at three or more plots are shown on the tri-plots. P-values from tests of significance of the
first and all canonical axes for each of the assemblages were: (a) 0.023, 0.028, (b) NS, 0.024, (c) 0.056, 0.033, (d) 0.000, 0.058, Formicidae NS,
NS, Coleoptera NS, NS, and were based on 9999 random permutations.
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334 331
c); and the exotic Acari: T. similis (E1) and Halotydeus
destructor (Tucker) (E2) (Fig. 4b and d). The CCA tri-
plot for epigaeic Collembola (Fig. 4c) further
separated the moderate, high and lime treatment plots
based on the negatively correlated variables pH and
organic carbon. Moderate treatment plots, with lower
pH and higher organic carbon, were characterised by
native Collembola: Australotomurus sp. (N16), Lepi-
docyrtus sp. (N8) and Katianninae sp. (N1), with the
above exotic Collembola dominating the high and
lime treatments. The dominance of exotic arthropods
in the higher treatment plots supports the trends
observed in Figs. 1 and 2.
5. Discussion
Despite the site’s history of low grazing pressure,
limited fertiliser application, and its apparent good
condition, considerable impact on the biodiversity of
the grassland had already taken place. This was
revealed most notably by the hypogaeic Acari where
90.8% of the 1747 sarcoptiform mites sampled from
the control plots were the exotic species T. similis.
Exotic species, although less dominant, were also a
significant component of the epigaeic Acari and
Collembola sampled from the control plots.
How different are arthropod faunas of putatively
good condition grasslands compared with grasslands
which have not been managed for production?
Within the production landscapes of south-eastern
Australia very few locations have escaped continuous
grazing by domestic livestock. King and Hutchinson
(1983) found that grazing reduced the number of
hypogaeic Collembola with average abundance over
the duration of an experiment; 10,600, 7900 and
4350 individuals m�2 for ungrazed, lightly grazed
and heavily grazed treatments, respectively. These
results support a history of light grazing for the
study site (average abundance in control plots was
7520 individuals m�2), but also suggest that Collem-
bola abundance may have been reduced and composi-
tion altered as a result (King and Hutchinson, 1983).
The average mite density in the two control plots was
considerably higher than King and Hutchinson (1983)
numbers demonstrating that a history of light grazing
and occasional fertiliser application had led to
significant increases in hypogaeic Acari abundance,
the community composition being dominated by a
single exotic species.
After only 4 years of fertiliser treatment and
grazing on these plots, exotic arthropods became more
abundant and some native species were largely
restricted to the control or low input treatments.
The environmental gradients responsible for these
differences included ground cover, exotic biomass,
pH, phosphorus, organic carbon and potassium, with
the first four variables significantly affected by the
treatments. Further, the variables: electrical conduc-
tivity, aluminium, calcium and sulfur, were omitted
from the analyses due to colinearity. Gradients in these
omitted variables may have also been responsible for
the differences in assemblages observed among plots.
Plots that received higher input management were
therefore characterised by higher P, EC, Ca, and S, and
higher exotic biomass and ground cover, and lower
organic carbon and Al compared with the control and
low treatment plots. In addition, the lime treatment
resulted in a large increase in pH.
Considerable work has been completed overseas on
the impacts of changed soil pH on soil-active
arthropods, both in terms of soil acidification due to
fertilisers and acid rain, and soil neutralisation with
lime (Hagvar and Abrahamsen, 1980; Hagvar and
Amundsen, 1981; Hagvar and Kjøndal, 1981; Hagvar,
1984). These studies have also shown that artificial
changes of soil pH can result in large changes in the
abundance and composition of Acari and Collembola
communities. In fact, Huhta et al. (1983) showed that
several published effects of fertilisers on the soil fauna
may have actually been related to changes in soil pH
rather than direct effects of the chemicals themselves.
Altering the pH of soils will result in changes of the
arthropod fauna, whereas the implications of liming
for biodiversity conservation in Australian soils are
less clear. Where long-term fertiliser application and/
or the sowing of legumes has resulted in soil
acidification (King and Hutchinson, 1989; Lockwood
et al., 2003), increasing pH with lime may well have a
positive influence on a site’s native biota by returning
the soil to a more ‘‘natural’’ state. However, many
Australian soils are inherently acidic (Scott et al.,
2000), including those studied in this project. Liming
soils with naturally low pH may lead to local
extinction of endemic acidophilic species. Recom-
mendations for broad-scale application of lime to
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334332
naturally acidic soils, therefore, need to be treated with
caution if biodiversity loss is not to be exacerbated.
Considerable heterogeneity was revealed between
assemblages sampled from the two sewage plots. In all
cases, one sewage plot was located on the CCA tri-
plots near the ‘‘higher’’ input treatments, and the other
was positioned near the control and low input
treatments. We expect that these differences in
assemblages are a direct result of different starting
and finishing levels of phosphorus and pH in these two
plots. By the completion of the experiment in 2002,
phosphorus in one sewage plot had only increased
from 9.7 to 24 mg kg�1 (150%), and pH had
commenced at 4.6 and dropped to 4.5. In the second
sewage plot (located near the high-input plots on
the CCA tri-plots) phosphorus had increased by
nearly 450% from 6.4 to 35 mg kg�1, and pH had
commenced at 4.9 and dropped to 4.75. These
differences in starting and finishing soil conditions
were most likely related to differences in soil type
within the two plots. The former plot was on yellow
Chromosol and Leptic Rudosol soils, while the latter
was on yellow Sodosol and Hydrosol soils.
Despite these differences in starting conditions, it is
possible that the results for these two plots represent
either side of a biological threshold for arthropods.
That is, a 150% increase in phosphorus resulting from
the application of sewage ash appeared to have little
impact on arthropod biodiversity, whereas, a 450%
increase had a dramatic impact in line with high inputs
of superphosphate and lime. To the authors’ knowl-
edge no studies have been published on the impacts of
sewage ash on native species. However, several
investigations have explored the ecological impacts
of sewage sludge application but most have investi-
gated its use in, and effects on, forest plantations
(Costantini et al., 1995; Crohn, 1995; Loch et al.,
1995; Bramryd, 2001), but see Yeates (1995), Larsen
et al. (1996), Andres (1999) and Cole et al. (2001)
for reported impacts on arthropods. Further experi-
mentation of rates of application of sewage ash and
resultant impacts on arthropod biodiversity are clearly
warranted.
The sensitivity of native grasslands to domestic
grazing and fertiliser use was a critical finding of the
study. Our results revealed a significant historical
impact of low-input management and a further
decline in native arthropod biodiversity through
reduced abundance and/or local extinction of native
species and increased dominance of exotic species
with increased fertiliser use and grazing intensity.
These results are in accordance with Australian
studies conducted on the northern tablelands of New
South Wales (King and Hutchinson, 1983; King
et al., 1985). Consequently, our study, together with
those of others, suggests that improving native
grasslands with fertiliser in the naturally nutrient
poor landscapes of south-eastern Australia may not
be ecologically sustainable from the point of view of
biodiversity conservation (see Groves et al., 2003).
However, given the state of development and the
economic importance of production landscapes in
south-eastern Australia, we do not expect that our
study will initiate major landuse change. Our
findings do, however, support the critical need to
identify, de-stock, protect and restore a representa-
tive network of native grassland ecosystems within
this highly productive agricultural zone of Australia
(Eddy, 2002). Our results also suggest that monitor-
ing attempts to restore degraded grassland systems,
as well as monitoring the impacts of sustainable
grazing management practices, may well benefit
from the inclusion of an arthropod monitoring
component (Paoletti, 1999).
Acknowledgments
This project would not have been possible without
the existence of the experimental plots which were
established in 1997. We thank other members of the
project team responsible for the original experiment
including; Colin Langford, Peter Simpson, Colin
Shields and Tony Ciavarella, and the Murray-Darling
Basin Commission for funding the original experi-
ment. The authors also thank Sarah McGeoch, Rob Le
Lievre and Colin Shields for efficiently taking care of
the field component of this project, and Kithsiri
Dassanayake for exporting the botanical data from the
original experiment’s database. Mark Dangerfield,
Simon Ferrier, Dieter Hochuli, Warren Mason, Chris
Nadolny, Lance Wilke and Brian Wilson provided
constructive comments on earlier versions of this
paper. This project was funded by the NSW
Biodiversity Strategy and the then NSW Department
of Land and Water Conservation.
I. Oliver et al. / Agriculture, Ecosystems and Environment 109 (2005) 323–334 333
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