J. Appl. Entomol.

The impact of ants on mineral soil properties and processes at

different spatial scales
E. L. H. Cammeraat1 & A. C. Risch2
1 Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, Nieuwe Achtergracht, Amsterdam, The Netherlands
2 Swiss Federal Institute for Forest, Snow and Landscape Research, Research Unit Community Ecology, Zuercherstrasse, Birmensdorf, Switzerland

Keywords
ecosystem engineers, infiltration, nutrient
cycling, overland flow, scale, soil quality
Correspondence
E. L. H. Cammeraat (corresponding author),
Institute for Biodiversity and Ecosystem
Dynamics, University of Amsterdam, Nieuwe
Achtergracht 166, 1018 WV Amsterdam,
The Netherlands.
E-mail: lcammera@science.uva.nl
Received: June 18, 2007; accepted: January
17, 2008.
doi: 10.1111/j.1439-0418.2008.01281.x
Abstract
Soil dwelling ants are important soil engineers that have a large impact
on the soil ecosystem. This is reflected in the alteration of soil properties
by ants due to burrowing activities, the accumulation of organic matter
and other nutrients in the soil, which, in turn, alters soil physical,
chemical and (micro) biological processes. A review of recent literature
on these fine scale soil changes resulting from ant activity is presented
in this paper. However, ants do not only affect soil properties at the fine
scale but also have effects at the hillslope and catchment scales with
respect to surface hydrological processes and ecosystem functioning. At
the finest scale, most studies showed changed soil textures, lowered bulk
densities and increased infiltration where ant burrows or mineral
mounds are prominent. Only a few studies investigated the effects of
ants on soil properties such as soil structure and aggregate stability,
which were improved by ant activity. Nutrients were generally consid-
ered to be concentrated in nests, and increased nutrient cycling involv-
ing microbial processes were reported. On a broader scale, the
information on nest densities and patterns is crucial to understand the
impacts of ants, but this information is scarcely available. It has been
reported that the activity of ants increases the heterogeneity of soil infil-
tration rates and nutrient concentrations at the landscape scale, which
in turn has consequences for the redistribution of sediments and nutri-
ents by geomorphological and biotic processes. A hierarchical descrip-
tive model focusing on dry land ecosystems is presented to incorporate
small-scale ant activity into broader scale hillslope and catchment pro-
cesses. However, this model has to be tested and more research is
needed, especially on these broader scale aspects, to be able to fully
understand the role of ants on the ecosystem and landscape scale.

carbon, nutrients and soil microbes within their

Introduction
Ants belong to the periodic endopedonic fauna (Hole
1981) and are present in almost all terrestrial ecosys-
tems (Hölldobler and Wilson 1990; Bolton 1994).
They have an important impact on soil physical,
chemical and biological properties by creating
macro-voids, galleries, chambers, organo-mineral soil
aggregates and by changing the composition of
nests (Lavelle 1997; Lobry de Bruin 1999; Seybold
et al. 1999; Karlen et al. 2003). The ancient Greeks
were well aware of the importance of soil digging by
ants as elucidated by the myth of the giant gold
digging ants (Herodotus 2007).
Ants are perceived as ecosystem engineers (Jones
et al. 1994; Folgarait 1998; Lavelle and Spain 2001),
but their importance in contributing to soil turnover

J. Appl. Entomol. 132 (2008) 285–294 ª 2008 The Authors

Journal compilation ª 2008 Blackwell Verlag, Berlin 285
The impact of ants on mineral soil properties
or bioturbation depends on both the type of ecosys-
tem and the ant species (Folgarait 1998). For exam-
ple 0.42 Mg/ha soil per hectare were turned over by
ants in a semi-arid ecosystem in Australia (Briese
1982), 10 Mg/ha were reported for soils in the wet
tropical and temperate zones from various ecosys-
tems (Paton et al. 1995). However, despite their
potentially large impact on soil properties, ants still
receive less attention than other macro-faunistic taxa
such as earthworms or termites (e.g. Léonard et al.
2004; Six et al. 2004). Yet, in geo-ecosystems, where
earthworms and termites are not predominantly
present, ants are among the most important soil
engineers, in particular in many sub-humid and
semi-arid areas.
This paper reviews the impact of ants as ecosystem
engineers within a geo-ecosystem response frame-
work and its consequences for soil quality and land
degradation. First, an overview will be given on the
impact of soil dwelling ants on fine scale soil proper-
ties (pedon scale) with emphasis on soil physical,
chemical and biological aspects. Secondly, we will
discuss the effects ants could have on soil properties
on ecosystem/landscape scales, by incorporating the
available information into a hierarchical model based
on the work of O’Neill et al. (1986).
Ants and Soil Properties: Fine Scale Effects
Effects on soil physical and hydrological properties
Fragment size, texture and aggregates
The burrowing activity of ants is well known
(Hölldobler and Wilson 1990; Paton et al. 1995;
Folgarait 1998). These insects pick up sand grains,
bite off fragments of soil aggregates with their man-
dibles (Hole 1981) and relocate this material. The
size of the soil particles transported by ants depends
on the size of the mandibles (Dostál et al. 2005),
and thus on the ant species. Ants have been shown
to form organo-mineral soil aggregates by excreting
chemical substances, which, in turn, alter the soil
structure and soil aggregate stability (Lavelle 1997).
For example, Cammeraat et al. (2002) found a sig-
nificant increase of soil structural stability of the
topsoil as a result of ant activity in semi-arid range-
land in Spain, but not for the subsoil of Messor
bouvieri nests. Decaëns et al. (2002) studied the
influence of soil biota, including ants, on soil bio-
genic structures and soil aggregates brought to the
soil surface. They found that especially ants of the
species Trachymyrmex sp. and Camponotus sp. contrib-
uted significantly to aggregates brought to the soil
286
E. L. H. Cammeraat and A. C. Risch
surface in Columbia, even though their nests occu-
pied less than 1% of total soil area. However,
despite these two studies that investigated the effect
of ants on soil aggregate stability, Six et al. (2004)
concluded in their literature review that the effects
of ants on such processes are not very well studied
and more research is needed.
Further, ants bring soil material from deeper soil
horizons close to or on top of the surface. This leads
to intensive mixing of soil material from different
soil depths and therefore changes soil textural prop-
erties (Paton et al. 1995; Folgarait 1998; review in
table 1). As many weathered soils have higher clay
contents with increasing depth, ant affected topsoils
commonly have a greater proportion of fine material
than non-affected topsoils. Several authors reported,
for example, increased level of silts and/or clays in
the mounds built by different ant species compared
to the surrounding soil confirming that the exca-
vated material is finer textured (MacMahon et al.
2000; Cammeraat et al. 2002; Whitford 2002; Dostal
et al. 2005; Azcárate and Peco 2007; Leal et al. 2007;
table 1). We are aware of only one study that
reported increased sand levels in the top of the
mounds constructed by Iridomyrmex greensladei on a
vertisol type of soil in New South Wales in Australia
(Nkem et al. 2000). Their contrasting result might be
explained by the fact that the smectitic clays in these
soils are sensitive to dispersion. Consequently, this
would increase the sensitivity of clay particles to ero-
sion, decrease mound stability, and lead to a selec-
tive enrichment of coarse particles. It should also be
kept in mind that besides the direct effect of ants on
soil textural properties, also indirect effects such as
overland water flow processes and rain-wash (Paton
et al. 1995) might influence the composition of the
excavated material found around ant nests and min-
eral nest mounds.
Pore space and bulk density
It is generally known that the excavation of soil
material by ants increases the (macro) porosity of
the soil, which in turn alters the ratio between the
soil solid and air phase and leads to lower bulk den-
sities. Most publications reviewed in our literature
study confirm such decreases (Lobry de Bruin and
Conacher 1990; : various ant species; Dean and
Yeaton 1993; : Messor capensis; Lal 1988; : various ant
species; MacMahon et al. 2000; : Pogonomyrmex sp.).
Although some other authors found no difference
between bulk densities from nests of Pogonomyrmex
rugosus and the surrounding controls in extremely
sandy soils (Wagner and Jones 2004; Wagner et al.
J. Appl. Entomol. 132 (2008) 285–294 ª 2008 The Authors
Journal compilation ª 2008 Blackwell Verlag, Berlin
 Table 1 Overview of significant effects of ants on various chemical and physical soil properties

Other soil Other

Soil physical macro-nutrients/ Microbial
Location Ant species Treatment Texture moisture properties pH* Org C Ntot, (NO)3), (NH)4+ P Remarks activity Reference

New Mexico, USA Pogonomyrmex barbatus, o ), o [NO3, NH4] + + [K, Mg] + + Wagner et al. (1997)
nest-control 6.4 [Ca] o
E. L. H. Cammeraat and A. C. Risch

Eastern Australia Aphaenogaster barbigula + (s) + (s) [Ntot]+ (s [Ca,Mg] +, [Na] o, [K]) (s) Eldridge and Myers
mound-control 5.8 o (n) [Ntot]o (n [K, Ca,Mg] + (n (1998)
(s= surface soil) o (n) [Na] o
(n= sub-surface soil) 5.2

Journal compilation ª 2008 Blackwell Verlag, Berlin

Netherlands Lasius flavus ) [bulkdens] - + o [Ntot, NO3] + [NH4] o + Blomqvist et al. (2000)
mound-control

J. Appl. Entomol. 132 (2008) 285–294 ª 2008 The Authors

N,S and C. America Pogonomyrmex sp. >Silt/ sand + [bulkdens] - + [Ntot, NO3, NH4] + + [K] + +/) MacMahon et al. (2000),
nest-control. (review)
New South Wales, Iridomyrmex <Clay [Spec. volume] o o, + [NO3, NH4] + + [K] o Nkem et al. (2000)
Australia greensladei >Sand 6.3 [Na,Ca,Mg, ESP] ) [EC]+
mound- control
Hessen, Germany Myrmica scabrinodis, o o, + Dauber and Wolters
Lasius niger, Lasius flavus 5.2 [Cmin]+ (2000)
Mound-control
Hessen, Germany Myrmica scabrinodis, ), 5.9 + [Nmin]+ + Dauber et al. (2001)
Lasius niger, Lasius flavus [Cbiom] +
mound-control
Almeria, Spain Messor bouvieri >Clay + [inf]+/ ) ), + [NO3,NH4] + + [K,Mg,EC] + Cammeraat et al. (2002)
nest-control [wr]+ 8.1
[struct]+
Alabama, USA Solenopsis invicta + [Ntot, NO3, NH4] + Depth Bender and Woods
mund-control (2003)
California, USA Messor andrei – ), + [Ntot]+ + + Boulton et al. (2003)
7.2
Amazonia, Atta sexdens [NO3]+, [NH4] o [Nmin,Nnit] o Verchot et al. (2003)
Brazil
Czech Rep., Lasius niger +, o + [Na, K] + Frouz et al. (2003)
Germany Mound-control 5.9 [Ca] o
and Slowakia spatial
Nevada, USA, Pogonomyrmex rugosus Wagner and Jones
Boulder City nest-control Clay + o [bulkdens] o ), 8.7 + [Ntot, NO3, NH4] + + (2004)
Kane Springs Clay) o ) ), 8.2 + + +
The impact of ants on mineral soil properties

287
 Table 1 Continued

288
Other soil Other
Soil physical macro-nutrients/ Microbial
Location Ant species Treatment Texture moisture properties pH* Org C Ntot, (NO)3), (NH)4+ P Remarks activity Reference

New Mexico, USA Pogonomyrmex rugosus o [bulkdens] o ), + [Ntot, NH4] + + time Wagner et al. (2004)
nest-control 7.1)7.5
Slovakia Lasius flavus >Silt [bulkdens] + +, – [Ntot]- + [K] + [C/N] + Dostál et al. (2005)
mound-control 4.2 [Ca, Mg] )
depth
Louisiana, USA Solenopsis invicta 7.4 + [K ] + Lafleur et al. (2005)
greenhouse [Na,Ca,Mg] o
texture depend.
The impact of ants on mineral soil properties

Louisiana, USA Solenopsis invicta – [bulkdens] ) o, [NO3] +,[NH4] o + [Na,K,Ca,Mg] +

Forest nest-control 4.3)4.5 o in plantations
spatial
Illinois, USA Formica montana +fio [bulkdens] + fi o +fio [Ntot, NO3, NH4] + fi o [DON] + fi o Lane and BassiRad (2005)
Acantha myops time
Czech Republic Lasius niger + ) [Ntot] ) + + Holec and Frouz (2006)
Lasius flavus [respir]o
Mound-control
Neotropics Atta laevitigata o [NO3 ]o,[NH4] +/o Agriculture Jiménez and Decaëns (2006)
Colombia Atta Landolti o [NO3]o,[NH4] + savanna,
Trachymyrmex sp. o [NO3] o,[NH4] + Gallery forest
nest-control
Nevada,USA Pogonomyrmex rugosus + [NO3,NH4] + + [Ndenit] + [respir]+ Jones and Wagner (2006)
nest-control
Nevada,USA Pogonomyrmex rugosus + + [NO3,NH4] + [decomp] + Wagner et al. (2006)
P. barbatus
nest-control
Richmond, USA No information ) ), [NO3, NH4] + [C min] ) Amador and Görres (2007)
6.6 [C biom] )
Spain Messor barbarus >Silt + [Ntot] + [K] + Azcárate and Peco (2007)
midden-control (acid)
NE, Brazil 18 species mixed >Clay [penetrability] + o, o [Ntot] o o [Ca, Mg, CEC]+ Leal et al. (2007)
6.9
New Mexico, USA Pogonomyrmex rugosus o o/+ Spatial Zaragoza et al. (2007)
Myrmecocystus depilis + o
Aphaenogaster cockereli o

+: significant increase, ): significant decrease in nest/mound with respect to control soil, o: no significant difference, + fi o: significant increase changes over time in no significant difference.
*pH value of the control soil. Org C = organic carbon or organic matter, bulkdens = bulk density, inf = infiltration, wr = water repellency, struct = soil structural stability, Nmin = N ineralisation,
Nnit = N nitrification, Ndenit = N denitrification, C min = C mineralisation, C biom = microbial biomass, decomp = decomposition, respir = respiration, spatial = spatial information available,
time = temporal change studied, depth = depth gradients studied, CEC is cat-ion exchange, DON = dissolved organic nitrogin, EC = electrical conductivity, ESP = exchangeable sodium percentage.

J. Appl. Entomol. 132 (2008) 285–294 ª 2008 The Authors

Journal compilation ª 2008 Blackwell Verlag, Berlin
E. L. H. Cammeraat and A. C. Risch
E. L. H. Cammeraat and A. C. Risch
2004), they did not provide any explanations for
their findings.
Infiltration rates
Higher infiltration rates are commonly reported in
ant affected areas compared to areas with no ant
activity (Lobry de Bruin and Conacher 1990;
Eldridge 1993; Cerdà and Jurgensen 2008). A clear
relationship has been detected between the size of
the ant nest entrance and infiltration rates (Eldrigde
1993) similarly to what Léonard et al. (2004) have
found during field- and in-silico experimental work
on termites in the Sahel. Wang et al. (1996) speci-
fied that only open nest entrances of Lasius neoniger
nests contributed to enhanced infiltration in Sparta
soils located in the lower Wisconsin river valley
(USA), and that many nest entrances were closed
during rainfall events, re-opening within 2 h after
the event. Eldridge and Myers (1998) suggested that
increased infiltration rates in nest entrances of
Aphaenogaster barbigula also facilitated the entrap-
ment of plant leaves, an important food resource,
that are transported by overland flow. In contrast to
these studies, Sarr et al. (2001) found that ants did
not significantly affect infiltration rates in the Sene-
gal and another study showed that some ant species
(Acromyrmex landolti) even constructed turrets at the
nest entrance to prevent water infiltration into their
nests (Navarro and Jaffe 1985). Further, Cammeraat
et al. (2002) showed that infiltration rates can be
affected by hydrophobic behaviour of the topsoil
around the nests of Messor bouvieri due to accumula-
tion of chaff and water repellent residues of plants
around, and in the nests. This resulted in lower infil-
tration rates in ant nests under dry conditions com-
pared to the surrounding non-ant affected soil,
whereas under wet conditions the ant nests showed
higher infiltration rates.
Soil moisture
The moisture status of mineral soil ant nests in rela-
tion to their surrounding soil is quite variable.
Wagner et al. (1997, 2004) and Zaragoza et al.
(2007) indicated no significant differences in soil
moisture in nests of Pogonomyrmex barbatus in dry
areas of the southwestern United States, whereas
Wagner et al. (2006) found higher values in similar
types of environments and the same ant species
(table 1). Also other studies detected no differences
in soil moisture inside and outside the nests (Mac-
Mahon et al. 2000; : Pogonomyrmex sp.; Cammeraat
et al. 2002; : Messor bouvieri; Table 1). Woodell and
King (1991) in Blomqvist et al. (2000) who worked
J. Appl. Entomol. 132 (2008) 285–294 ª 2008 The Authors
Journal compilation ª 2008 Blackwell Verlag, Berlin
The impact of ants on mineral soil properties
in the Netherlands suggested that the lower soil
moisture contents that they found in ant mounds
were related to increased evaporation as more pores,
burrows and channels were present in mounds.
From this, one may expect that ant nests show
stronger moisture dynamics with increased infiltra-
tion and evaporation rates compared to non-ant
affected soils. Furthermore, water infiltration is also
related to the extension of the vertical galleries and
tunnels and the size of the nests (MacMahon et al.
2000), factors that likely are nest age and ant species
dependent.
Conclusion
Our review shows that soil physical and hydrological
properties seem to strongly depend on the ant species
involved, the type of soil and the ecosystem inhab-
ited as well as on the initial conditions of the soil.
Even within the same ecosystem and soil type and
the same ant species under study, different effects on
soil physical or hydrological properties have been
described. In general, ants either have no effect or
increase clay contents in the surface soil and increase
infiltration rates. Ants have rarely if ever been found
to increase sand content or decrease infiltration rates.
Effects on soil chemical properties
The bulk of information available on the effect of
ants on soil chemical properties is related to organic
carbon (Corg), pH and (different forms of) nitrogen
(N), while only few studies focused on how ants
alter the availability of other soil nutrients. Gener-
ally, the observed differences in Corg and nutrient
levels are attributed to the accumulation of food
(leaves, seeds, ‘honeydew’) or excrements like saliva
and droppings in nest areas or separation and trans-
location of food and soil material during nest build-
ing (Eldridge and Myers 1998; Folgarait 1998;
Lavelle and Spain 2001; Frouz et al. 2003 and many
others). However, we would also like to point out
that nutrient levels, especially those related to
mobile and soluble organic substances [dissolved
organic carbon (DOC), dissolved organic nitrogen
(DON)], salts containing nitrate NO3), ammonium
NH4+, sodium (Na+), potassium (K+), magnesium
(Mg2+) and calcium (Ca2+), and phosphorus (P)
might also be affected by leaching processes in the
soil, especially in humid environments.
Carbon and nitrogen
Many authors reported higher Corg levels in semi-
permanent ant nests (see reviews by Lobry de Bruyn
289
The impact of ants on mineral soil properties
and Conacher 1990; Folgarait 1998; MacMahon
et al. 2000; table 1). Frouz et al. (2003) and Holec
and Frouz (2006) studied mound building ants
(Lasius niger, Lasius flavus) and found, however, no
significant differences for Corg and Lane and Bassiri-
Rad (2005) showed that differences between
mounds and controls became smaller with increasing
ant colony age.
Nitrogen levels are commonly differentiated in
NO3), NH4+ or DON. Generally, the concentrations
of these forms of nitrogen show higher or
unchanged levels in ant nests compared to the con-
trol soils (table 1). The reason is likely related to
changes in microbial activity and N turnover of
organic material or to excrements added to the nest
by ants (e.g. MacMahon et al. 2000; Jiménez and
Decaëns 2006; Jones and Wagner 2006; table 1).
Nests of various species of harvester ants in dry eco-
systems showed increased levels for both NH4+ and
NO3- including higher turnover rates, whereas ant
nests and mineral mounds from the tropics and
more humid environments showed a larger diversity
in N related properties and turnover rates. For exam-
ple, Verchot et al. (2003), studying Atta sexdens in
Brazil found no differences in NH4+, but clearly
higher levels of NO3- in the subsoil around the nests.
They also found hardly any differences in minerali-
zation and nitrification rates between control soil
and the nests and concluded that N-cycling was not
increased in the nests. Similar findings were reported
by Lafleur et al. (2005). In contrast, higher ammo-
nium concentrations have recently been found with
no increase in nitrate (Jiménez and Decaëns 2006),
no change in the N pool (Leal et al. 2007) or even
decreased N levels (Holec et al. 2006). They indicate
that ants could have quite variable effects on soil C
and N pools and cycling. It should also be kept in
mind that these processes are strongly related to
other environmental factors such as precipitation
regimes or soil moisture. Especially in dry environ-
ments differences in soil moisture may affect the
decomposition and respiration rates (Wagner and
Jones 2004; Jones and Wagner 2006). Also the activ-
ity and age of ant nests is important regarding their
role as nutrient accumulation and recycling hotspots,
shown by an initial increase followed by a decrease
of nutrient accumulation in the ant nest as observed
by Lane and BassiRad (2005).
pH
It generally has been suggested that ant activity low-
ers soil pH in alkaline soil nests and increases soil pH
in acidic soil nests (Petal 1980 in Folgarait 1998).
290
E. L. H. Cammeraat and A. C. Risch
Our literature review on newer studies could, how-
ever, only confirm this for alkaline soils (Cammeraat
et al. 2002; Boulton et al. 2003; Wagner and Jones
2004; Wagner et al. 2004; Lafleur et al. 2005;
table 1). In these studies, the decrease in pH can be
explained by increased Corg levels. The decomposi-
tion of the Corg leads to the production of organic
acids, which then leads to a decrease in pH. This pre-
sumed relationship is not clear from the data pre-
sented in the literature on neutral or acid soils,
where no effects, increases or decreases in soil pH
were observed (table 1).
P, Na, K, Ca and Mg
The presence of higher concentrations of P in ant
nests and mineral ant mounds has been observed by
most authors (in 15 of 16 studies in giving details on
P; table 1). Only the recent publication of Leal et al.
(2007), who studied a mixed group of 18 ant species
in Brazil, found no significant difference in P
between nests and the control soil. However, most
of the publications do only provide information on
total, but not on available P (e.g. P-Olson). Espe-
cially in Ca-rich or weathered soils with high
amounts of Fe- and Al-oxides and in allophanic
soils, available and not total P is limiting for vegeta-
tion and a differentiation in P-types should be made
to enable conclusions with regard to nutrient avail-
ability in ant nests.
Sodium, K, Ca and Mg are related to soluble salts
and show various results (table 1). The most con-
stant finding is that K is generally higher in ant nests
or mineral mounds compared to the surrounding soil
(Wagner et al. 1997; MacMahon et al. 2000; Cam-
meraat et al. 2002; Frouz et al. 2003; Dostál et al.
2005; Lafleur et al. 2005; Azcarate and Peco 2007;
table 1). Only, the study of Eldridge and Myers
(1998), who worked with nests of Aphaenogaster bar-
bigula, showed lower K levels in the surface soil of
nests. However, in the subsoil of the nest they also
detected higher K levels compared to the control,
whereas Nkem et al. (2000) (Iridomyrmex greensladei)
reported no significant differences at all.
Conclusions
Again, the information provided in the literature
showed highly variable effects of ants on soil chemi-
cal properties and nutrient turnover rates, which
generally depended on the ecosystem type and ant
species under study. Further, ants seem to have a
clear acidifying affect when located in alkaline soils
and generally tend to increase N cycling and P con-
centrations.
J. Appl. Entomol. 132 (2008) 285–294 ª 2008 The Authors
Journal compilation ª 2008 Blackwell Verlag, Berlin
E. L. H. Cammeraat and A. C. Risch
Effects on soil biological properties
Bardgett et al. (2001) discussed the general impor-
tance of the soil fauna in transporting materials,
their influence on soil water movements as well as
their impact on soil biophysical properties. Although
several types of soil macro-fauna contribute to
changes of soil properties, their effects are very dif-
ferent. Jouquet et al. (2006) differentiated between
animals with permanent and ephemeral nests or
burrows and classified them as ‘extended phenotypic
engineers’ such as ants and ‘accidental engineers’
such as earthworms. Spatio-temporal heterogeneity
in landscapes is strongly increased by the activity of
‘extended phenotypic engineers’ whereas ‘accidental
engineers’ homogenize the soil through bio-turba-
tion processes. Recently, an increased number of
papers have been published giving attention to these
‘extended phenotypic engineers’ and showed that
ants affect the decomposition and microbial activity
in belowground nests and mineral mounds (e.g.
Dauber et al. 2001; Verchot et al. 2003; Wagner
et al. 2004; Wagner and Jones 2006; Holec et al.
2006; Amador and Görres 2007; table 1). Generally,
higher amounts of microfauna and –flora have been
detected in ant nests (Wagner et al. 1997; Dauber
and Wolters 2000; MacMahon et al. 2000; Boulton
et al. 2003; Holec and Frouz 2006; Amador and
Görres 2007; table 1).
Ants and spatial patterns: effects at hillslope and
landscape scales
Although it might be an over-simplification, ant
nests, as shown in our review on fine scale effects,
function as local sinks of nutrients, water and other
resources. However, the enrichment of Corg, nutrients
and microbes in ant nests also depends on landscape
position as shown by Whitford and DiMarco (1995)
working with harvester ants (Pogonomyrmex rugosus)
in the Chihuahuan Desert, USA. These authors
reported that ant nests located in slope bottom areas
did not show marked differences in these properties
when compared to surrounding soils. They explained
these results by the fact that ant nests located at the
slope bottoms were subject to disturbances by flood-
ing and sediment deposition. On the hillslopes, how-
ever, they found larger and significant differences
between nests and surrounding soil. The spatial pat-
terns of soil physical, chemical and hydrological prop-
erties created by ant nests are therefore not only
important at the scale of the ant nest or mound, but
have also important consequences for pedological,
J. Appl. Entomol. 132 (2008) 285–294 ª 2008 The Authors
Journal compilation ª 2008 Blackwell Verlag, Berlin
The impact of ants on mineral soil properties
geomorphological and hydrological processes at hills-
lope or catchment scales. This has also been reported
by Cammeraat et al. (2002) who showed that infiltra-
tion was significantly higher under wet than dry
hydrophilic conditions in ant nest areas of Messor bou-
vieri. Further, it has been shown that the pattern and
density of ant nests affects the accumulation or dis-
persal of resources throughout entire ecosystems.
When hillslope scale ant nest cover is dense, the ant
nests act as important buffers for water, sediments
and water-transported nutrients (Cerdà and Jurgen-
sen 2008), in particular in bare and sparsely vegetated
environments. This, in turn, will increase the hillslope
scale heterogeneity with nutrient hotspots and locally
depleted areas in-between the hotspots. Under condi-
tions with limited ant nest occurrence and sparse veg-
etation cover, surface water flow will increase,
causing erosion, depleting the topsoil. This combina-
tion of direct alteration of the soil by soil dwelling
ants and its consequences for geomorphological pro-
cesses may affect the pedological and hydrological
processes and development. We summarized the
potential impact that ant nests could have at different
spatial and temporal scales in fig. 1 based on a hierar-
chical model originally proposed by O’Neill et al.
(1986, 1991) and adapted to geo-ecosystem processes
by Cammeraat (2002). Erosion processes and hydro-
logical leakiness are also strongly influenced by vege-
tation cover in general and by vegetation patterns in
particular (Bochet et al. 1999; Valentin et al. 1999;
Ludwig et al. 2002, 2005). However, to include the
effects ants would have on the vegetation would go
beyond the scope of this review.
Unfortunately, there are relatively few studies on
the spatial distribution and density of ant nests and
foraging holes at large scales. Moreira et al. (2004)
illustrated the spatial pattern of nest and foraging
entrances for three ant colonies (Atta laevigata) and
found 60 entrances for one nest. Warburg and
Steinberger (1997) studied the spatial pattern of two
types of ants of the genus Messor spec., and concluded
that the distribution of ant nests can be considered
as random. Holec et al. (2006) investigated the effect
of vegetation patch pattern on the distribution of
Lasius niger nests in the Czech Republic. They
reported quite dense patterns with 165 active nests
on 2500 m2; small nests were aggregated whereas
medium and large nests were randomly distributed
and occurred more often in less vegetated areas.
Finally, Wagner et al. (2004) showed that the nest
surface area of Pogonomyrmex barbatus increased over
time following a power function reaching a maxi-
mum after approximately 13 years, with a decline in
291
The impact of ants on mineral soil properties
Fig. 1 Hierarchical geo-ecosystem model, showing harvester ant related relationships between fine, intermediate and broad scale processes and
geo-ecosystem properties for a dry land environment where bare areas are abundant.
densities thereafter. Overall, the ant nest densities
ranged from 0.1–10% cover of the soil surface (e.g.
Eldridge 1993; Cammeraat et al. 2002; Jones and
Wagner 2004; Lane and BassiriRad 2005). To under-
stand the importance and impact of ant nests on
ecosystem properties at different scales, more infor-
mation is needed on size, cover and spatial distribu-
tion. Only by integrating research on ants, soil and
surface properties (including vegetation) across hills-
lope and catchment scales, we will be able to under-
stand these complex geo-ecosystem processes, which
have important implications for remediation mea-
sures in degrading areas, for agricultural systems and
for soil quality maintenance and improvement.
Acknowledgements
The unknown reviewers of the manuscript are
thanked for their very constructive comments,
which improved the paper substantially. The orga-
nizers of the 2007 EGU session on ‘Ants in the Soil
System. A hydrological, chemical and biological
approach’ are thanked for stimulating to write this
review.
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