Agriculture, Ecosystems and Environment 74 (1999) 425–441

Ants as bioindicators of soil function in rural environments

L.A. Lobry de Bruyn ∗
School of Rural Science and Natural Resources, Ecosystem Management, University of New England, Armidale, NSW, 2351, Australia

Abstract
Farmers are being encouraged to redress the problems of land degradation by implementing a series of changes in their land
management. However, there are few tools to monitor soil condition which can reliably inform farmers of the state of their soil.
This paper examines ants and their importance in ecosystem functioning, principally their ability to maintain or restore soil
quality in rural environments, and their value as bioindicators of soil quality. The reader is provided with a summary of those
studies which have recorded the species composition and activity of ant fauna in agricultural soils and naturally vegetated
environments. However, it is rare to find studies which sample both environments concurrently. An overview of the types
of measures available to assess ant biodiversity is followed by a discussion of the current status of biopedological research
into ant–soil relationships and the trends and gaps in current research knowledge, principally in the areas of water and soil
movement and nutrient effects. Lastly the paper seeks to forecast the future directions of ant biopedological research, and to
identify the type of research required to fully appreciate the value of ants as soil quality indicators to monitor the sustainability
of the soil resource. Ants should be considered an important group of soil fauna, yet they have received meagre attention in
comparison with other soil fauna such as earthworms. There is a need for specific experiments to test the hypothesis that ants
can be used as bioindicators of soil quality. ©1999 Elsevier Science B.V. All rights reserved.
Keywords: Soil quality; Ants; Bioindicators; Soil macrofauna; Agriculture; Biodiversity; Biopedological

1. Introduction soils, in comparison with naturally vegetated en-

This review of ants in rural environments concen-
trates on their importance in ecosystem functioning,
principally in their ability to maintain or restore soil
quality. The biology and ecology of ants is not consid-
ered in detail here and for this information the author-
itative work by Hölldobler and Wilson (1990) should
be consulted. The objectives of this review are:
1. To examine the need for soil sustainability indi-
cators and their desirable attributes.
2. To establish an understanding of the ant fauna
that exists, survives and proliferates in agricultural
∗ Tel.: +61-2-6773-3119; fax : +61-2-6773-2769
E-mail address: llobryde@metz.une.edu.au (L.A. Lobry de Bruyn)
vironments.
3. To overview the types of measures available to
assess ant biodiversity.
4. To establish the current status of biopedological
research into ant–soil relationships and to examine
the trends and gaps in current research knowledge.
5. To forecast future directions in ant biopedological
research, including the value of ants as soil quality
indicators to monitor the sustainability of the soil
resource.
The approach has followed two courses of action. In
the process of writing another paper (Lobry de Bruyn,
1997) the author asked eminent Australian researchers
in agricultural ecology questions pertaining to the di-
rection of soil macrofauna research, the trends and

0167-8809/99/$ – see front matter ©1999 Elsevier Science B.V. All rights reserved.
PII: S 0 1 6 7 - 8 8 0 9 ( 9 9 ) 0 0 0 4 7 - X
426 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
gaps in current research, and the importance of soil
macrofauna as a tool to monitor soil sustainability. The
other avenue was to examine Current ContentsTM over
1993–1998 for published research in this area; in ad-
dition a range of review articles summarising research
before 1993 was consulted.
2. The need for ants as bioindicators of soil
function and their desirable attributes
The rapid rate at which land (0.3–0.5% of arable
land per annum) is being degraded by erosion, salin-
isation, and contamination (Lal and Stewart, 1992)
is the most compelling reason to establish bioindica-
tors of soil function. The interactions of soil quality
along with soil stability and soil resilience contribute
to the sustainable use of the soil resource (Lal, 1993).
Soil quality is defined by Doran and Parkin (1994) as
“The capacity of the soil to function within ecosystem
boundaries to sustain biological productivity, maintain
environmental quality, and promote plant and animal
health.”
Soil macrofauna, which includes ants, can be used
as an integrative measure of soil quality, assuming
their importance in regulating soil processes which are
vital to the continued formation of soil and as pro-
tection against soil degradation as highlighted in sev-
eral international workshops on sustainable agriculture
(Hawksworth, 1991; Doran et al., 1994).
These bioindicators are required to monitor changes
in soil quality and to provide early warning of adverse
trends and identify problem areas (Pieri et al., 1995).
Governments around the world (The Standing Com-
mittee on Agriculture, 1991; Acton and Gregorich,
1995; Munasinghe and Shearer, 1995) have identified
three areas of agricultural sustainability which need
to be monitored to ascertain whether the agricultural
management practices that are being advocated are
achieving the goal of sustainable agriculture. These are
the natural resource base, economic viability of agri-
cultural production, and other ecosystems which are
influenced by agricultural activities. The protection of
the natural resource base—soil—and the prevention of
its further degradation are critical to the development
of sustainable agriculture and continued production of
food and fibre (Acton and Gregorich, 1995). Govern-
ment reports (Hamblin, 1992; SCARM, 1993) have
also identified soil biota as potential bioindicators of
soil sustainability at the farm level, although presently
there is little rigorous experimentation to test their
validity in agroecosystems (Pankhurst et al., 1995;
Doran et al., 1996).
An additional reason for examining soil biota,
specifically ants, and their relationship with soil qual-
ity and soil sustainability is that farmers need indica-
tors of soil quality which they can easily and reliably
use to monitor soil sustainability. Ease and reliability
are significant properties, as farmers are unlikely to
adopt the soil sustainability indicators derived by sci-
entists if they require too much technical expertise,
expense and time to monitor, and produce results that
are difficult to interpret (Baker and Dalby, 1994).
Farmers have been slow to adopt sustainable manage-
ment practices because they cannot see the benefits
of the new techniques and perceive a higher risk and
uncertainty with them (Vanclay, 1992; Carter, 1994).
At the same time there is a strong desire by farmers
to monitor their farm goals and determine if changes
in farm management are leading towards a more sus-
tainable farming system. A simple monitoring tool
which illustrates trends in soil quality may act as a
persuader to change or affirm management practices
(Romig et al., 1995).
Ants are not often considered as soil biota, and in-
deed are sometimes excluded in definitions of soil
biota (e.g. Killham, 1994; Wood, 1996). In relation to
their interaction with the soil, soil biota can be cate-
gorised into three groups: epigeic (those which process
organic matter on or near the soil surface), endogeic
(those living in the soil) and anecic (those which trans-
fer materials between soil and litter habitats) (Bouché,
1977). Ants could be placed in either the anecic or
epigeic groups. Soil biota are also categorised accord-
ing to size, and ants are considered to be macrofauna
(body width 2–20 mm). It seems that ants and their
role in soil functioning are often overlooked in favour
of other invertebrate groups such as earthworms or
termites. The author maintains that ants have a re-
silient role in maintaining soil quality because of their
ability to survive in agricultural soils despite the va-
garies of climate and disturbance regimes, albeit as a
less diverse fauna. Moreover, Majer (1983), stated that
ants conform to most of the following attributes which
are considered desirable qualities of an indicator. The
indicator should be:
 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
• closely related to one or more of the assessment
goals,
• important to the overall structure and function of
the agroecosystem,
• responsive to a range of environmental stresses,
• easily measured and quantifiable,
• easily interpreted,
• holistic and
• have integrative effects over time (Pankhurst, 1994).
Others have been more cautious in their endorse-
ment of bioindicators and suggest that much more ba-
sic research is required (Paoletti and Bressen, 1996).
There is an important impediment in using ant bio-
diversity (measured simply by species richness) as an
indicator of a healthy soil. Firstly, there is a need to
understand and be able to identify which species or
groups of species have key functions in the mainte-
nance of energy and material flows through an ecosys-
tem (Silver et al., 1996). It has been assumed that a
soil ecosystem with low biodiversity is less resilient,
more vulnerable to perturbations, and ultimately not as
able to function as well as a soil ecosystem with high
biodiveristy. However, little is known about the con-
tribution of individual species or groups of species to
ecosystem functioning and the effect of their removal
from the soil ecosystem. There is a tendency in ant
biopedological research to emphasize a single species
or groups of species. These ant species have been se-
lected because of their dominance either in terms of
foraging activity, visibility of nest sites or abundance
as gauged by nest density. However, the justification
for concentrating on a single species is rarely given,
and few studies have quantified the impact of the total
ant community. Establishing which ant species are im-
portant in terms of soil quality requires an understand-
ing and quantification of their impact on the soil pro-
file, and their association with soil types.
3. Ant biodiversity in rural and naturally
vegetated environments
World-wide, the largest numbers of ant species
are found in Australia and in the Tropics (Table 1).
Understanding of ant biodiversity in modified land-
scapes, especially agricultural, is fragmented. Invari-
ably, research into ant biodiversity has focused on
natural environments or those affected by mining or
427
Table 1
Number of described soil species of Formicidaea
Region Number of species
Neotropical Region 2233
Australia 1100
North America (+ USA) 585
USA 400+
France 180
Sweden 61
Denmark 49
Finland 47
Norway 46
UK 46
New Zealand 23
a Source: Groombridge (1992).
forestry (Majer, 1984, 1985; Majer et al., 1987) ei-
ther at the species or community level. Earlier studies
of the biodiversity of soil biota in agricultural soils
(Abbott and Parker, 1980) usually have sorted the in-
vertebrates to a higher taxonomic level (class, order or
family level) or have used morphospecies (Oliver and
Beattie, 1993). Again, few studies traverse ecological
boundaries by examining ant biodiversity in both rural
and naturally vegetated environments. Tables 2 and 3
summarise data on ant biodiversity in rural environ-
ments and adjacent naturally vegetated environments.
Many studies have found high species diversity in
natural environments within Australia (Majer, 1978;
Greenslade and Mott, 1979; Andersen, 1983). The
studies presented in Table 2 indicate that species
richness is nearly always lower in farmland environ-
ments than in naturally vegetated areas. However, the
information from each study can rarely be directly
compared either because of differences in soil type,
rainfall or intensity of agricultural activity (Table 2).
Work by Lavelle and Pashanasi (1989) in a range
of land management types (forest, traditional and
high input cropping systems, traditional and improved
pasture and palm-trees) in the tropics (Peruvian
Amazonia) recorded ant biomass between 0.1 and
0.8 g f wt m−2 . The proportion of biomass represented
by ants in comparison with other soil macrofauna (ter-
mites and earthworms) was minor (0.02–5%) (Lavelle
and Pashanasi, 1989). However, in terms of popula-
tion density, ants make a more significant contribution
(7–53%) when compared with the total abundance
of macrofauna in each land management type, espe-
cially in traditional cropping (53%) and traditional
pastures (49%). According to a study of Petersen
428 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441

Table 2
Number of ant species found in rural and neighbouring naturally vegetated habitats
Location Rural environments Naturally vegetated Reference

Species Habitat Species Habitat

number number

Chihuahuan Desert, USA – – 3–36 Desert grass/shrubland Whitford, 1996

Kellerberrin, Australia 32 Wheat/Pasture rotation 65 Heath shrubland Lobry de Bruyn, 1993b
Kellerberrin, Australia 29 Wheat/Pasture rotation 89 Wandoo woodland Lobry de Bruyn, 1993b
Katanning, Australia 19 Improved pastures, grazed – – Twigg et al., 1982
Deniliquin, Australia 27 Native pastures, grazed – – Briese and Macauley, 1977
Wubi-Perenjori, Australia 12 Lupin stubble 35 Open woodland Keals and Majer, 1991
Emmet Vale Australia 17 Native pastures, grazed 22 Saltbush Briese and Macauley, 1977
Kellerberrin, Australia – – 15–40 Gimlet woodland Abensperg-Traun et al.,
(unpublished)
Biebrza, Poland 1–6 Meadows on peat soils 39 Pine forests Petal, 1991
Costa Rica, South America 23 Coffee plantations – – Perfecto and Snelling, 1995

(1982), the calculated biomass of ants in Europe
ranged from 1.9 g day wt m−2 for three ant species
at Mols, Denmark, to 12 g day wt m−2 for Formica
rufa in Siberian forests. Although the ant biomass
was much lower for grassland sites in America and
Africa (0.1–0.5 g day wt m−2 ; Petersen, 1982), ants
are nevertheless considered important relative to
other soil fauna in these environments. More recent
work by Petal and Kusinska (1994) in Poland gave
ranges of ant biomass for Myrmica spp. and Lasius
niger as 0.8–6.3 g day wt m−2 . In Australia, Davison
(1987) found that the colony biomass of Chelaner
whitei ranged from 2.24–161.05 g day w m−2 , to
6.21–55.27 g day wt m−2 for C. rothsteini.
The advent of agriculture usually produces a de-
cline in ant biodiversity. There are three common ex-
planations for the loss of species in agricultural envi-
ronments: the changes in microclimate conditions ad-
versely affect development of larvae and pupae (Höll-
dobler and Wilson, 1990), foraging activity and nest-
ing places; food availability is reduced from use of
agrochemicals (fertilizer, herbicides and pesticides)
and reduction in litter and soil organic matter; and
interactions with other species leads to alterations in
community structure. Agricultural soils are then sub-
ject to varying levels of cultivation which causes dis-
ruption to the habitat and nesting places of ants, but the
effects are often transitory (Lobry de Bruyn, 1993b).
However, the observed patterns may also be indepen-
dent of the above factors and merely arise as an arte-
fact of site selection.
Agricultural systems which are considered to be
more sustainable are those which imitate those envi-
ronmental conditions found in natural environments.
Brussaard et al. (1988) outlined a new research di-
rection for agroecosystems which would compare the
functioning of a soil–crop ecosystem in conventional
and integrated farming systems. The outcome would
be an agroecosystem which promotes better soil struc-
ture, improved nutrient use efficiency and controlled
diseases, pests and weeds more effectively. They fore-
saw changes in agricultural practices that would re-
duce nutrient inputs, soil tillage and use of biocides
(Brussaard et al., 1988). The general consensus is that
ant biodiversity and abundance increase under mini-
mum tillage and stubble retention. A study by Rad-
ford et al. (1995) found four times the density of ant
colonies under a no tillage system compared with a
conventional tillage system. Wang et al. (1996a) found
more ant burrow openings in the less irrigated and no
tillage plots than in the more ploughed and irrigated
treatments. However, the previous year had shown no
effect of agricultural practices. In a study of the ef-
fects of soil type and cultural practices on the fire ant
(Solenopsis invicta) in South Louisiana sugar cane-
fields, Ali et al. (1986) recorded the highest densi-
ties in fields with clay soil and high levels of ground
cover. The ant nests of Messor spp. were more abun-
dant in unploughed than in ploughed plots, and the
shrublands showed higher numbers of nests than the
adjacent agricultural fields (Diaz, 1991). An exam-
ple from a study in Kellerberrin in Western Australia
L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441 429
430 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
shows that the advent of agricultural development has
produced a decrease in ant species richness by as much
as 50% (Lobry de Bruyn, 1993b). However, ant activ-
ity was similar (measured by foraging activity and ac-
tive nest density) in farmland and naturally vegetated
land of the same soil type. Such similarity provides
hope for the further development of sustainable agri-
cultural systems, because it is through their activities
of nest building and foraging that ants have the po-
tential for abating soil degradation. The environmen-
tal factors which control ant species distributions were
most highly correlated to climatic factors, soil type
and vegetation structure and floristics (Hölldobler and
Wilson, 1990).
4. Methods to estimate ant biodiversity
4.1. Techniques to sample ants
Techniques available to sample ant fauna include
baits, nest mapping, hand sampling, litter and/or soil
extraction, and pitfall traps (Edwards, 1991; Paoletti
et al., 1991). The choice of sampling technique is
very much dependent on the aim of the survey. For a
complete census of the ant biodiversity of an area it
is advisable to undertake a combination of the above
techniques. Majer (1997a) and others (Adis, 1979;
Abensperg-Traun and Steven, 1995), have critiqued
the efficacy of relying on pitfall trapping as the only
sampling procedure to collect ants; concerns can be
summarised as follows:
• The diameter of the pitfall trap affects the num-
ber of species caught (Abensperg-Traun and Steven,
1995).
• Some ant species are easier to trap than others
(Marsh, 1984).
• The true representation of the total ant fauna, espe-
cially in habitats with dense litter cover and cryptic
or rare species (Andersen, 1991), is unlikely.
• The technique only samples surface-active ants.
Andersen (1983, 1991) also noted that the close
proximity of a pitfall trap to an active nest or forag-
ing trail can lead to a higher abundance of certain
species or, in the case of species which have low
locomotor activity from its nest, will lead to an
underestimation of its abundance.
431
• The technique gives no indication of absolute inver-
tebrate density.
• The technique has difficulty in establishing whether
the species caught is residing in the habitat sampled
or only foraging.
Despite the above caveats, there are some obvious
advantages to pitfall traps: they are simple to use, op-
erate continuously through day and night, and yield
high numbers of ants which then can be statistically
analysed (Majer, 1997a). Greenslade and Greenslade
(1971) and Greenslade (1973) have examined in
greater detail some of the operational concerns. More
consideration needs to be given to foraging distances
of ants, which vary considerably, from less than 0.5 m
up to 50 m (Hölldobler and Wilson, 1990). This is
especially important for examining variation in ant
activity in agricultural plots which are in close prox-
imity to each other, to ensure the independence of
plots. Abensperg-Traun and Steven (1995) concluded
that the optimal relationship between trap diameter
and trap density will be a compromise between cap-
ture efficiency (experimental design), logistics and
the use of complementary sampling methods.
Lobry de Bruyn (1993a, 1993b) combined pitfall
trapping with the mapping of ant nests to give a more
accurate representation of the ant fauna, especially
those species with a more active role in soil func-
tion. Crist and Wiens (1996) assessed the spatial pat-
terns of prominent ant nests using aerial photography,
vector-based Geographic Information Systems (GIS)
and ground-truthing. This method is only appropriate
for conspicuous nests. The density and dispersion of
soil ant nest activity of less conspicuous nests can be
mapped using fixed quadrats over a smaller area, e.g.,
5 m2 (Wang et al., 1996a) or 6.25 m2 (Lobry de Bruyn,
1993b). Furthermore, mapping has to be carried out
over a year to cover the seasonal fluctuations in species
activity and to monitor the activity of individual nests.
Nest monitoring is important for defining the spatial
and temporal impact of a particular ant species whose
nests may have been identified as having a beneficial
effect on water infiltration or to assess the influence
of ant activity on soil turnover rates.
Another sampling procedure is to use soil cores,
which will give an absolute population density. The
standardised sampling method designed by the Trop-
ical Soil Biology and Fertility (TSBP) programme
relies on 10 samples of 25 cm × 25 cm × 30 cm soil
432 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441

monoliths, taken 5 m apart, and hand-sorted into

four strata (litter, 0–10, 10–20, 20–30 cm) (Ander-
son and Ingram, 1988). In Australia, Robertson et
al. (1994) and Radford et al. (1995) took 10 samples
of 18 cm × 18 cm × 15 cm soil monoliths to quantify
macrofauna density in agricultural plots, but recorded
low ant abundances. Soil cores or monoliths are con-
sidered inappropriate for estimating the contribution
of ants to soil ecosystem functioning, and are more
suitable to less mobile soil macrofauna such as earth-
worms. Fig. 1. A drawing of an Aphaenogaster sp. which is a soil specialist
and a major contributor to soil turnover in semi-arid environments
To achieve a comprehensive understanding of ant
of Australia. Scale bar = 1 mm.
activities and their effects on soil properties there has
to be an appreciation of the temporal and spatial ex-
tent of their activity, which can be gained through pit-
fall trapping and mapping as well as other techniques
(Lobry de Bruyn, 1993a; Perfecto and Snelling, 1995).

4.2. Classification of ants

To reduce the complexity of ant taxonomy but still

develop an interpretable model for examining ant com-
munity structure and its relationship with land use,
Andersen (1986, 1990, 1993) has further developed a
classification of ants into functional groups that was
originally proposed by Greenslade (1978). This classi-
fication has gained wide acceptance, especially in Aus-
tralia, but still suffers from problems of interpretation.
Andersen (1997) has recently adapted the functional
group classification for North American ant communi-
ties. The functional group classification is based on the
adaptations employed by other ant genera in avoiding
interaction with Iridomyrmex spp. which are extremely
abundant, active and aggressive in arid environments
in Australia. The characteristics of each group can be
summarised as follows:
1. Dominant species – Iridomyrmex spp., which are
highly abundant and aggressive, influence the dis-
tribution and activity of other ant species.
2. Subordinate species – Usually Camponotus spp.
which because of their size, periods of activity
and submissive behaviour are able to coexist with
Iridomyrmex spp.
3. Climate and soil specialists – These ant species
(Melophorus spp., Meranoplus spp., Aphaenogaster
spp. (Fig. 1) and Notoncus spp.) reduce their
competition with the dominant species by mor-
Fig. 2. A drawing of Rhytidoponera metallica which is an op-
portunist and is often found in disturbed environments such as
agricultural soils. Scale bar = 1 mm.
phological, physiological and/or behavioural
adaptations.
4. Cryptic species – These ant species live, feed
and forage within the soil and litter (Brachypon-
era lutea), thus avoiding contact with other ant
species.
5. Opportunists – These ant species (Rhytidopon-
era spp. (Fig. 2), Paratrechina spp.) are scav-
engers and predators, poor competitors and are of-
ten found in disturbed habitats such as farmland.
6. Generalised Myrmicinae spp. – These include ant
species of the genera Monomorium, Cremato-
gaster and Pheidole (Fig. 3) which are flexible in
their foraging times and feeding requirements.
7. Large, solitary foragers – Large ant species of
the genera Myrmecia, Bothroponera and Odon-
tomachus are solitary predators that do not inter-
act greatly with other ants.
Taxonomy needs to be developed in order to im-
prove our understanding of how soil communities
function. The work of Oliver and Beattie (1993, 1996)
suggested that the most cost-effective and accurate
assessment of species turnover would be to allow
 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
Fig. 3. A drawing of a Pheidole sp. (major and minor) which
is found in high nest densities in agricultural soils in semi-arid
Australia, and builds nests to depths of 60 cm. They have flexible
foraging times and feeding requirements. Scale bar = 1 mm.
non-specialists to sort specimens into morphospecies
and to target sampling to known optimum periods of
activity.
5. Current status of biopedological research into
ant–soil relationships: trends and gaps
There is little understanding of the role that ants
play in soil processes and plant growth, especially the
mechanisms by which ants exert their influence on soil
processes, in particular soil structure, soil hydraulic
properties and nutrient cycling. There has been no at-
tempt to quantify the relationship between ant activity
and pasture production or crop yields. Research find-
ings often present the end results of the interaction be-
tween ants and soil processes rather than information
on how ants facilitate, impede or have inconsequential
influences on soil function.
5.1. Water movement
The connection between ant activity and changes
in soil physical properties has been given meagre at-
tention, with research showing that the channels and
galleries of ants may either impede or facilitate water
infiltration (Lobry de Bruyn and Conacher, 1990). It
433
might be expected that ant channels contribute to in-
creased aeration, drainage and improved root growth,
given that biopores greater than 1 mm in diameter have
been found to be responsible for such benefits (Oades,
1984). However, there is little evidence to confirm this
phenomenon.
Ants may facilitate water movement by increasing
porosity. For instance, Denning et al. (1977) found
that 16% of the volume of the upper section of a
Formica cinerea mound (140 cm across and 40 cm
high) contained macropores with a saturated conduc-
tivity of 10 000–44 000 cm/day. This is up to 4000
times the conductivity needed to cope with the max-
imum precipitation expected in a 24 h period. Majer
et al. (1987) found that the mean infiltration rate on
ant nests (8.3 ± 5.4 min/l) was much faster than on un-
affected soil (27.2 ± 3.9 min/l). In areas of semi-arid
Australia under natural vegetation, studies have been
conducted to assess the influence of ant nests on water
infiltration (Eldridge, 1993). These studies recorded
ponded water infiltration rates four times faster over
Aphaenogaster barbigula nest entrances (5–43 mm)
than on the control soil. On the other hand, the soil
around some ant burrows has been found to be bare.
Thorp (1967), for example, described a 2–10 m circle
of bare ground around the nest of a harvester ant on
the Great Plains of North America, which could im-
pede water infiltration and encourage soil erosion.
However, few studies have directly assessed the in-
fluence of ant biopores on water infiltration (Eldridge,
1993; Lobry de Bruyn and Conacher, 1994a; Wang et
al., 1996b). The density and form of these biopores
needs to be assessed in situ (Lobry de Bruyn and
Conacher, 1994a), using a disc permeameter (Perroux
and White, 1988).
Lobry de Bruyn and Conacher (1994a) examined
the influence of ant biopores on water infiltration un-
der negative tensions (−5, −10, −40 mm) and positive
pressure (+10 mm) in the wheatbelt of Western Aus-
tralia. Ant biopores were found to conduct water four
to six times faster than the control soil, but only under
ponded conditions, which would occur under high en-
ergy rainfall events. The studies by Wang et al. (1995,
1996a, 1996b) demonstrate the importance of quanti-
fying the impacts of ants, in this case a single species
– Lasius neoniger – on bioporosity and water flow
by examining the spatial and temporal patterns of ant
burrows, nest architecture and water infiltration down
434 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
ant burrows. All the measurements were conducted in
agricultural fields.
The effectiveness of ant biopores in conducting wa-
ter under ponded conditions is dependent on several
factors. The ant nest biopores must remain open dur-
ing a rainfall event which eventuates in overland flow,
in order to absorb runoff which could otherwise lead
to erosion. During heavy rainfall events there is a
likelihood of raindrop impact or turbulent overland
flow dislodging the surrounding ant nest spoil and
sealing the nest entrance, as was observed in Dur-
rokoppin farmland after a rainfall event of 73 mm in
24 h; it was not clear at what point during the rain-
fall event the sealing occurred (Lobry de Bruyn and
Conacher, 1994a). Wang et al. (1996b) found the main
reason why ant burrows were not significant in wa-
ter flow was that they tended to close during an irri-
gation or rainfall event. It was found that after only
5 mm of irrigation, 80% of the burrows had closed,
but within 2 h the equivalent proportion had reopened
(Wang et al., 1996b).
Wang et al. (1995) examined the nest structure of
Lasius neoniger by impregnating the nests with dental
gypsum. The nest volume varied from 20 to 250 cm3 ,
with the galleries concentrated in the top 30 cm
(100.33 cm3 ), and only 15.35 cm3 below 30 cm to a
depth of 70 cm. Other studies have used lead (Markin,
1964), latex (Brian and Downing, 1958), and wax
(Salinitri, 1976) to characterise ant nest morphology.
Lobry de Bruyn, (1990) found polyurethane foam to
be very effective in making a cast of an ant nest, es-
pecially those with entrance holes greater than 2 mm
in diameter. Ant nests may extend to depths of up to
0.8 m (Gray, 1974), but in some cases can be quite
shallow (0.2 m) (Ettershank, 1968; Eldridge, 1993).
In addition to their capacity to remain open to con-
duct water, the spatial distribution and density of ant
nest biopores are also important factors that influence
water drainage. Table 3 shows various studies which
have mapped the density of ant nests and/or burrows
in agricultural or naturally vegetated environments,
but rarely concurrently. On average, ant nest openings
rarely exceed 3 per m2 in farmland sites. For effective
infiltration of water, it might be assumed that ant nest
openings need to be sufficiently evenly spaced and
dense, but often ant nests are highly aggregated, espe-
cially if they are connected underground to each other,
which is often the case in ant species which are poly-
calic. Nevertheless, Sisson and Wierenga (1981) ob-
served that a large fraction of the water passes through
a small fraction of the plot area, because of the spatial
variability of the steady-state infiltration rate. The ag-
gregated distribution of ant nests may not, therefore,
be a problem. Indeed, the localised, higher densities
may prove to be more efficient in intercepting and ab-
sorbing overland flow than a more evenly distributed
pattern of nests with lower densities. Finally, the sea-
sonal fluctuation of nest densities influences the pe-
riod during which ant nests would be effective in con-
trolling water erosion. Ant nests will probably remain
open only until farmers begin to plough; afterward, it
is doubtful whether the biopores would be of much
importance until the ants reopen their nests, usually a
month after seed is drilled.
As mentioned above, ant biopores have a positive in-
fluence on steady-state water infiltration rates, but only
under ponded conditions (at positive heads). Thus, the
frequency and duration of ponded conditions in farm-
land needs to be investigated. It would be necessary
to carry out rainfall simulation experiments on repli-
cated plots with and without ant nests, and compare
the time to ponding and volume of runoff generated
from the two treatments.
5.2. Nutrient effects
The impact of ants on nutrient cycling is not ex-
tensively documented in rural environments (Lobry de
Bruyn and Conacher, 1990). In recent years ecologists
identified the ant mounds of Pogonomyrmex barbatus
(Wagner et al., 1997) and Formica podzolica (Lesica
and Kannowski, 1998) in naturally vegetated environ-
ments as contributing positively to the resource base
for other biota, and nest mounds of Lasius flavus, La-
sius alienus and Formica rufibarbis to resource patch-
iness (Dean et al., 1997). Researchers have examined
the chemical composition of nest spoils and com-
pared them with adjacent soils which are unaffected
by ant activity. Typically, researchers have examined
ant species with conspicuous nests, such as Beattie
and Culver (1983) and Culver and Beattie (1983),
who examined nests of Formica canadensis in Col-
orado, USA. Such conspicuous, long-lived nests are
usually recorded at low nest densities (5/ha; Frouz
et al., 1997). Epigeic invertebrates, such as ants, de-
 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
posit undecomposed organic material at the soil sur-
face or in chambers within the soil. The decomposed
organic material may move down through the soil ei-
ther by biological mixing or by water and gravity, or
be dispersed over the soil surface by overland flow
or wind action. Rogers and Lavigne (1974), Wiken
et al. (1976) and Levan and Stone (1983) studied the
influence of ants on the chemical properties of soils
and, in general, found some nutrient enrichment at ant
nest sites. Presumably these nutrients are redistributed
when the nest is eroded. This may occur quickly with
ant nest structures which are unstable, not long-lived
and not compacted. Such nutrient enrichment is point
centered and its overall influence depends on the den-
sity of ant nests. Ant-modified soil invariably con-
tains increased levels of nutrients such as organic C,
PO4 and N, but sometimes the increases are small
or negligible. However, organic matter content is not
always greater at ant nests compared with adjacent
soils. Greenslade (1974), for example, found no ap-
preciable change in organic matter content between
Iridomyrmex purpureus mounds (5.47%) and the ad-
jacent soil (5.5%). Petal (1978, 1980) and Petal and
Kusinska (1994) investigated the environmental condi-
tions in ant nests to determine if conditions for organic
matter mineralisation and its humification are more ad-
vantageous in the ant nest than in the surrounding soil.
Their findings confirmed that total carbon content and
humic acids were lower in anthill soils than in soils
surrounding the ant nest, which indicates that condi-
tions in the nest were more favourable for organic mat-
ter mineralisation and less favourable for its humifi-
cation (Petal and Kusinska, 1994). Frouz et al. (1997)
found a higher decomposition rate of autochthonous
material, higher respiration activity and higher micro-
bial counts in the nests of Formica polyctena in a
spruce plantation than the surrounding soil. Wagner
(1997) also found that soil from ant nests had signifi-
cantly higher nitrogen mineralisation rates, and in an-
other study recorded 30-fold higher densities of mi-
croarthropods and 5-fold higher densities of protozoa
in soil from ant nests than in surrounding, control soils
(Wagner et al., 1997). Hence ant nests, especially those
which are conspicuous and long-lived, constitute an
important source of resource variability and availabil-
ity for other organisms.
The following ideas have been expressed to explain
alterations of nutrient status at the nest site. Levan and
435
Stone (1983) found that the immediate sub-mound soil
had elevated pH and levels of PO4 and Ca and lower
levels of organic matter than the corresponding control
soil. They argued that equal or lower organic matter
content compared with the surrounding soil reflected
the soil’s origin as surface soil/subsoil composites. A
simple mixing model can be used to explain elevated
nutrient levels but is inadequate to explain the extent
of nutrient increase. In cases where nutrient enrich-
ment exceeds the expected, Levan and Stone (1983)
suggested that inputs from exogenous sources must
account for such values. Briese (1982) claimed that
accumulation of C, PO4 and N at the nest site arose
from the decomposition of seeds and plant material
discarded at the nest entrance. Wiken et al. (1976) ex-
plained the increases in organic matter in ant-affected
sections of the soil pedon as the result of deposition
of somatic exudates, because microscope examination
did not reveal incorporation of forest litter or humified
material. Higher concentrations of these substances
(C, N and PO4 ) were maintained in the nests for at
least 2 years after the nests had been abandoned
(Czerwinski et al., 1971).
Lobry de Bruyn (1990) identified high percentages
of carbon at various depths down the nest systems of
two species of Rhytidoponera, whereas the other ant
species (Pheidole sp. 10) had higher percentages of
carbon in the top 2 cm than in the control soil. These ar-
eas of increased carbon could encourage plant growth,
and seeds located in nest middens were observed to
germinate. The spoils of most ant nests would also en-
courage germination because they are less compacted
than the control soil. It was further observed that newly
dug nests of Pheidole sp. D10 (most of which occurred
in farmland), did not show a significant increase in
carbon or other nutrients at a depth of 2 cm in compar-
ison with the control soil, whereas more established
nests of Pheidole sp. D10 in the naturally vegetated
sites did show an increase in carbon at the same depth
(Lobry de Bruyn, 1990). Hence, age of the nest, or
more appropriately the age of the nest midden, will
influence the degree to which soil chemical properties
are modified.
5.3. Soil movement
The movement of soil by ants (bioturbation) has im-
portant implications for soil formation rates and the
436 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
redistribution of soil particles, nutrients and organic
matter. The significance of ants to nutrient cycling is
closely related to the amount of soil they move during
nest construction and to the longevity of the nest site.
The basic technique used to construct nests by ants
is by pulling out soil particles with their mandibles,
and carrying a load of loosened soil to another place
where they deposit it. Hole (1981) described the exca-
vation activity of ants and termites as that of “miners
that seize sand grains or bite off fragments of aggre-
gates with their mandibles, leaving a pitted surface on
gallery walls”. This burrowing technique sharply con-
trasts with that of earthworms, which tunnel by push-
ing or eating their way through soil. Ants construct
nests to reduce the problems of weather variations,
to have a central base for the storage of food, and to
locate the queen and brood in a defensible structure
(Sudd and Franks, 1987).
The effects of a total ant community on the for-
mation of soil texture profiles requires an assessment
of bioturbation rates. The variables which affect the
depth and rate of ant bioturbation are: nest longevity,
which includes duration of habitation and growth and
erosion rates when the nests are active and senes-
cent; mass of soil held in ant spoils (excavated soil
brought to the soil surface), and nest densities. Table
3 shows that there have been few studies assess-
ing bioturbation rates by the total ant community
in semi-arid environments of the southern hemi-
sphere (Briese, 1982; Lobry de Bruyn and Conacher,
1994b). Humphreys (1981), Eldridge and Pickard
(1994), and Lobry de Bruyn and Conacher (1994b)
assessed the bioturbation rates of a single species
of Aphaenogaster in semi-arid environments, and
found it was responsible for significant movements of
soil (Table 3). The consequences of ant bioturbation
and its effects on the soil texture profile have varied
(Lobry de Bruyn and Conacher, 1990), ranging from
clay enrichment (Baxter and Hole, 1967; Salem and
Hole, 1968; Wiken et al., 1976), to no apprecia-
ble alteration in comparison with the surface soil
(Greenslade, 1974), to an increase in coarse material
(Mandel and Sorenson, 1982; Cowan et al., 1985).
Wang et al. (1995) estimated that Lasius neoniger
could turn over the upper 30 cm of soil once every
2800 years for a corn field, and once every 1000
years for the grass borders, where ant nest densities
are three times greater.
Determination of nest/spoil longevity is the most
problematic factor in estimating soil turnover rates
and making long-term predictions. No accurate tech-
nique for dating ant nests exists except by monitoring
nest growth from inception to final decay. Eldridge
and Pickard (1994) monitored the nest entrances of
Aphaenogaster barbigula in fixed quadrats for nearly
2 years, and found that nest entrances lasted 9 months,
and changed about twice a year. Reliable estimates of
spoil decay by erosion requires observations over pe-
riods longer than 10 years. The longevity of ants nests,
although variable, ranges from several months to about
10 years (Greenslade, 1974). With agricultural distur-
bance, nest longevity may be reduced. Lobry de Bruyn
(1993a) monitored ant nests in farmland over a crop-
ping season, and found that although nest entrances
disappeared after cultivation, they quickly reappeared
in less than 2 months.
The percentage surface area occupied by ant nests
is usually less than 1% of the soil surface, but occa-
sionally may be as high as 10% (Table 3). In Aube-
nas, France, which experiences a sub-mediterranean
climate, Aalders et al. (1989) estimated that 5% of
the ant spoil was transported by erosion and the re-
mainder was compacted and returned to the soil. The
end result of such processes is a gradual accumulation
of soil material on the soil surface by ant burrowing
activity.
6. Research implications
A priority for future research is to quantify the role
of ants in soil processes, and to elucidate their function
in agroecosystems. This objective would be realised
by integration of ant research with other related areas
and the inception of multidisciplinary projects (Faeth
et al., 1991 cited in Doran et al., 1996). There is an in-
creasing demand for real-time information on the con-
dition of the soil resource and for suitable indicators
which can monitor soil quality and act as early warn-
ing indicators of land degradation (De Soyza et al.,
1998).
The value of ants as indicators of soil quality to
monitor the sustainability of the soil is still unclear
(Linden et al., 1994; Lobry de Bruyn, 1997), even
though they have been used widely in establishing
the success of mining rehabilitation (Majer, 1997b).
 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
More research is required to understand the relation-
ships among ant community structure and particular
soil types, land use types and climatic regions in ru-
ral landscapes such as attempted by Way et al. (1997).
Giller et al. (1997) emphasised caution in establishing
causal links between soil biodiversity and sustained
soil fertility, and stressed that the scale of investiga-
tion can influence research findings. One of the major
obstacles to identify definitive links between ants and
soil processes is that the majority of research projects
have been observational. There has been little manip-
ulation or true experimentation of macrofauna such
as demonstrated in experiments carried out by Bohlen
et al. (1995) on earthworm populations in large-scale
field experiments in agroecosystems.
A greater understanding of the habitat needs of ants
in relation to treatment of the soil ecosystem is re-
quired. Even then, widely recognised observations of
population changes or species composition fluxes may
be a result of factors other than farm management.
Confounding factors such as geographic location or
timing of sampling are not always considered but can
significantly affect results. Ants may be useful indica-
tors at the farm scale, especially as part of a broader
system of ecosystem process indicators. Any advances
in this area will also simultaneously require the de-
velopment of taxonomic tools that could be used by
non-specialists to identify specimens.
Another limitation of ant biopedological research is
that funding agencies are concerned with agricultural
land use only and have a 3–5 year life span for projects.
This precludes studies of native fauna in naturally veg-
etated regions or on transcending boundaries between
agricultural and naturally vegetated land. The consid-
erable time frame required for this type of study also
constrains research outcomes. In Australia there have
been few studies which have examined ant communi-
ties for more than 1 or 2 years. Often conclusions have
been based on a sampling strategy with low intensity
and frequency accompanied by scanty or fragmented
information regarding environmental conditions. Ob-
viously, climatic variables will strongly influence the
outcome of the sampling activity.
Comparisons between research projects require a
standardised sampling protocol. Yet this would seem
impossible to achieve considering the diverse nature of
biopedological studies and the various logistical con-
straints. Ant sampling could be restricted to strategic
437
points in the year, with similar environmental condi-
tions.
Researchers have often utilised data from well es-
tablished experimental sites which were initiated for
other purposes, rather than setting up completely new
experiments. However, this precludes a baseline sur-
vey of the ant fauna, and hence does not consider the
patchiness of the population before experimentation.
Moreover, the experimental design usually is not ap-
propriate for examining biota which are mobile. When
experimental plots are long and narrow sampling is
limited to in situ methods such as soil cores or hand
sorting sods of soil. In these circumstances, pitfall
traps will not necessarily sample specimens which re-
side within the sample plot.
To test the importance of ants in nutrient cycling
and soil structure maintenance, one must compare
ant-inhabited soils to similar soils without ants. This
is difficult because the soil could have been influenced
by ants at any time during its formation. For examin-
ing ant activity the usual strategy has been to use areas
which show obvious signs of ant activity, such as an
ant nest, and compare them with areas where there is
no activity (Lobry de Bruyn, 1990; Holt et al., 1996).
An understanding of the limits of sustainable land
use requires identification of the critical biological in-
dicators of soil quality and their thresholds and hence
of soil sustainability (Lobry de Bruyn, 1997). Soil
resilience is an important determinant of the soil’s
response to degradative processes, and has been high-
lighted in the international scientific arena as an issue
of global concern, especially under current agricul-
tural land use practices (Lal, 1993; Greenland and Sz-
abolcs, 1994). There is a need to test the role of ants in
soil resilience and to define the critical limits of a soil
property from which the soil can recover to its initial
state, and the rate of that recovery. Lobry de Bruyn
(1997) emphasised the type of research direction
which is required if soil macrofauna are to be consid-
ered in maintaining a healthy, functioning soil. The
most pressing need is the commitment to develop rig-
orous, adequately-funded and long-term experiments
that measure soil biota at various levels (population or
community or biological process level), and to relate
these changes to other aspects of soil quality. Research
needs to take a holistic approach in order to understand
the function of ants in healthy and degraded soils.
Examining the relationship between ants and soil
438 L.A. Lobry de Bruyn / Agriculture, Ecosystems and Environment 74 (1999) 425–441
processes could lead to potential use of ants as a surro-
gate measure of soil processes essential to ecosystem
functioning (Nash et al., 1998). Ultimately bioindica-
tors of soil function should enable farmers to decide
if their resource is declining in quality/condition, is
stable or is in a process of renewal.
Acknowledgements
The author thanks colleagues who responded to the
questionnaire, anonymous referees, and Dr. Hamish
Sturrock for improvements to the manuscript.
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