ArticlePDF Available

Volume 2, Issue 2, December 2024 Population Genetics of the Meat Ant (Iridomyrmex purpureus) (A Literature Review)

Authors:
Derna University Journal of Medical Sciences
(Open Access Database-Peer Reviewed Journal)
Website: http://dujms.uod.edu.ly/
ISSN:3005-236X(Online), 3005-2351(Print)
Arab Impact Factor: 0.87

Volume 2, Issue 2, December 2024
11
Mahboba, 2024
Population Genetics of the Meat Ant
(I
ridomyrmex purpureus
)
(A Literature Review)
*Mahboba Aldareh
1Foundation Department, Faculty of Medical Technology, University of
Tobruk, Tobruk, Libya. Email: Mahboba aldareh @tu.edu.ly.
https://www.doi.org./10.58987/dujms.v2i2.2
Received: 20 10 2024 Accepted: 30 12 2024 © DUJMS 2024
ABSTRACT:
The meat ant (Iridomyrmex purpureus) is a native species of ant in Australia, exhibiting
extensive distribution across the eastern and southern regions of the nation. Population genetic
methodologies evaluate the genetic makeup of biological populations and the variability in
genetic components that arise due to the influence of diverse factors, including the
mechanisms of natural selection. Researchers in the field of population genetics strive to
achieve their aims by developing abstract mathematical models that describe gene frequency
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 12
dynamics, endeavoring to draw inferences from these models concerning the likely patterns of
genetic variation observed in real populations, and juxtaposing the findings with empirical
data. Due to climate change, ectotherms, particularly insects in temperate zones, face
significant thermal stress. Population genetics of meat ants residing in a temperate ecological
zone. Assessing critical thermal limits elucidates the effects of rising temperatures on
organisms. The meat ant served as a model organism for a comprehensive study of thermal
tolerance. The critical thermal maximum is influenced by the ramping rate in thermos limit
respirometry assays. Findings indicate that thermal responses in meat ants generally do not
differ among populations, with the exception of western inland populations, and climatic
factors and elevation showed no significant impact on thermal tolerance across these
populations. Because of studying genetic systems among insect populations offers insights
into genetic diversity, relatedness, and breeding strategies. In this article, I summarize
population genetics of meat ant (I. purpureus).
KEYWORDS: Australian Ants, Population Genetics; Causes of Population
Genetics, The Meat Ant (Iridomyrmex purpureus).
INTRODUCTION
Ants are arguably the most well-
known and ecologically remarkable
group of insects in the world, and
that is not because of their potential
as indicators of the health of
terrestrial ecosystem but it is also
because ants play an essential role
in the Australian environment
(Andersen, 2000). Ants also use the
most complex forms of chemical
communication of any creatures and
their social organization offers an
illuminating contrast to that of
human beings (Hölldobler &
Wilson, 1990). There are three
characteristics of ants ‘organization:
(1) cooperative brood care; (2) non-
reproductive workers or helpers; and
(3) overlapping adult generations
(Sudd & Franks, 1987). There are
also two more factors that make ants
so important: their value as objects
of biological study and their
importance in the economy of
nature (Wheeler, 1925). Ants are
species without hierarchical or
central control, they have the ability
to reach group level patterns and
they coordinate behaviour of
thousands of individuals (Detrain, et
al., 2001).
In rural environments, ants play
important roles in the ecosystem as
bioindicators of soil quality (Lobry
de Bruyn, 1999). Other ecological
functions of ants include
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 13
promoting seed germination as
seed dispersers, and ants also
interact with seeds of specific
plants and are responsible for
promoting some plant vegetative
growth (Wilson, 1990) and
contributing to reforestation of
many ecosystems (Peternelli, et al.
2004). In addition, ants play
essential roles in incorporating
organic matter and making it
fertile, and in soil aeration
(Moutinho, et al., 2003).
Ants, like all insects, have three
body parts; the head, the thorax, and
the abdomen. They also have two
compound eyes, two antennae and
maxillary palps. The antennae help
the ant to identify their mates and
enemies while the maxillary palps
help the ants to detect scent.
There are many diverse ant species,
but only 1,300 Australian species
have been scientifically identified
and named to that time (Andersen,
2002).
Ants are known to be social
insects as they live in colonies
(Bourke & Franks, 1995). The sizes
of the ant colonies depend on the
ant species. An ant colony consists
of a mother queen and her progeny;
the latter are mostly female ants
that are less fertile than the queen
or infertile, and daughter workers
(Sudd, 1967). The female ants
leave their colony to search for
food (Larson & Larson, 1967),
while the main role for workers is
to stay in the maternal nest and
support the queen so she can breed
more offspring (Sudd & Franks,
1987).
Most of the ant species live
within the ground. Their nest
entrances are open and may be
located under a rock or a log. In the
forest, many ants build their nests
by using different patterns; some
build their nests above ground by
utilizing trunks and hollows in the
branches of trees, and others create
their nest cavities via overlapping
leaves with each other by silk while
several ant species build their nests
in rotten wood or in the deep layer
of leaf litter (Burwell, 2007).
The ants ‘method of gathering
diverse foods depends whether they
are typically scavengers or
predators. The infertile female s
duty is to look for food for the
larvae. The larvae are only limited
to solid foods. The other ants,
especially the workers, are known
to be liquid feeders. This means
they gather sugary liquids such as
honeydew, sugary nectar and plant
secretions (Andersen, 2002).
Other ant species are more
specialized and have one mode of
feeding. Some are hunters of one
particular prey; some depend on
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 14
gathering seeds while others feed
on sugary liquids only. The
honeydew ants tend to create a
symbiotic relationship with the sap
sucking insects. They offer them
protection and in turn they share
their honeydew with the ants
(Shattuck, 2000)
Ants can be important
ecological engineers of habitats.
When they exist in large numbers
of colonies near another, they can
manipulate plant growth and
animal numbers.
Their relationship with plants
in which they extract floral nectars
while spreading pollen is a famous
example of mutualism (Sudd &
Franks, 1987).
Ants are a source of food for
many animals such as insects,
birds, mammals, and reptiles. Since
most ants live in the ground, they
help aerate the soil through their
excavations and tunnelling
(Greenslade, 1974).
Ants are known to help
spread plant species. Australia has
the largest plant species whose
seeds are spread by ants
(Gammanset et al., 2005). In
Australia, there are some different
species of ants shown below in
Table (1).
Table:(1). Some Different Species of Ants.
The spatial structure of a
population determines how natural
genetic difference is distributed
between and within populations, for
example, how population genetic
variations have differed from each
other and how much the populations
harbors genetic differentiation
(Seppä et al., 2009). In order to
survive, a population must be able to
adapt by altering its genetic structure
in response to the unavoidable
changes in the environment. It must
produce and reproduce new
phenotypes and genotypes improved
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 15
to the changing conditions (Mettler
& Gregg, 1969). There are some
factors that determine the changes in
population genetic structure.
Eliminated genetic differentiation
(by effective drift) and an introduced
new genetic generation (by fast
migration) determine the genetic
variation in the population; thus,
allele frequencies are diverged
among population by genetic drift,
while migration counteracts this
process. The essential assessment of
genetic population structure is not
only to understand species evolution
and the population generally, but
also to identify the population s
capability to keep their genetic
differentiation from being
compromised and changed (Seppä et
al., 2009).
Ants are haplodiploid organisms,
which mean the females emerge
from fertilized eggs, and males
from parthenogenesis; the two
occurrences of ploidy levels can be
utilized as an essential tool for
genetic analysis (Diehl, et al.
2002). Both the social behaviour
and genetic structure within and
among colonies are affected by
haplodiploid (Seppä et al., 2009).
Therefore, excellent opportunities
exist to investigate different
evolutionary and biologically
different populations using ant
populations and their various social
and highly complex colonies
(Antagalli, et al., 2013). Genetic
differentiation within and between
populations could be impacted by
the ants ‘social behaviour (Sugg, et
al., 1996). Several social insects
show differences in their social
behaviour that could possibly
impact their reproductive and
mating strategies; therefore, the
social insect provides a unique
opportunity to study population
genetic structure (Pamilo, et al.,
1997; Ross, 2001). In particular,
ant societies follow a special
strategy because they build their
nest as a sedentary and permanent
place (Rüppell et al., 2003)
In ecosystem function, ants
play role in terms of constituting a
great part of the animal biomass as
well as they present as ecosystem
engineers, and ants also are
important in regarding to ground
processes by the changes of the
physical and chemical environment
and through ants ‘impacts on
plants, soil organisms, and
microorganisms (Folgarait, 1998).
Due to the ants ‘abundance,
diversity, and long history, ants
constitute the important insects in
terrestrial ecosystem (Fernandez &
Delsinne, 2013).
1. Population genetics
Population genetic techniques
assess the genetic constituents of
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 16
biological populations and the
variation in genetic components
that are caused by the operation of
different factors, as well as natural
selection. Scientists in population
genetics pursue their objectives by
formulating abstract gene
frequency dynamics mathematical
models, trying to retrieve
conclusions from these models
regarding the probable patterns of
genetic disparity in actual
populations, and comparing the
results against experiential data
(Wright, 1969)
Studies in population genetics
also investigate the distribution of
allele frequency and variation is
influenced by the four major
evolutionary processes: mutation,
genetic drift, gene flow, and natural
selection. It similarly considers the
aspects of population structure,
recombination, and population
subdivision. Additionally, it tries to
clarify phenomena like speciation
and adaptation (Charlesworth, et
al., (2003). Many studies have
investigated the genetic
constituents of biological ant
populations and the variation in
genetic components that are caused
by different environmental factors.
Ants are social insects belonging to
the family Formicidae, and, along
with interrelated bees and wasps,
the order Hymenoptera (DeHeer &
Herbers, 2004). Ants evolved in the
mid- Cretaceous period from wasp-
like ancestors about 130 million
years ago and branched out after
the development of flowering
plants. Over 12,500 out of an
approximated total of 22,000
species have been categorized.
They are effortlessly recognized by
their unique node-like structure that
creates a slender waist and elbowed
antennae‖ (Wilson, 1990).
Colonies of ants can vary in
size from a small number of a
dozen predatory ants living in tiny
natural cavities to extremely
organized colonies that can inhabit
large territories and comprise
millions of ants. Larger colonies
consist generally of barren wingless
females creating castes of
"soldiers", "workers", or other
specialized groups. Some ant
colonies have a number of fertile
males (drones) and a number of
single fertile females (queens). The
colonies at times are depicted as
super-organisms since the ants
seem to operate as a cohesive
entity, jointly working as one to
support the colony (Hölldobler &
Wilson, 1990).
Ants have colonized nearly
every land habitat on Earth. The
only habitats lacking native ants are
Antarctica and a small number of
secluded or inhospitable islands.
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 17
Ants flourish in nearly all
ecosystems and can form 15-25% of
the Earth s animal biomass. Their
accomplishment in such numerous
environments has been credited to
their ability to transform habitats
and their social organization, tap
resources, and protect themselves
(Wilson,1990). Their lengthy
coevolution with many other
animals and plants has led to
parasitic mimetic, commensal, and
mutualistic relationships.
1.2 Causes of genetic
variation in ant populations
Once populations are
separated from each other, genetic
drift, the accumulation of novel
mutations, and differing selection
could drive genetic variations
between them (Tsutsui & Case,
2001). The homogenizing influence
of gene flow counters the genetic
variations (Slatkin, 1987). The
genetic variation in the population
is affected by several factors: how
fast migration produces new
differentiation to the population as
well as how effective drift
eliminates genetic variation. In
ants, specifically, the genetic
variation is influenced by the
genetic and social structure of the
colonies (Seppä et al., 2009) as the
social behaviour has an essential
impact on separating the genetic
differentiation within and between
populations (Sugg et al., 1996).
Therefore, sociality, migration, new
colony development, and genetic
variation are the main factors that
determine the change of population
genetics in ants.
1.2.1 Sociality
Sociality represents kin
cohorts as an extra hierarchical
level in populations. Due to insects
‘organization of life style and
ecological domination, social
insects have played a key role in
understanding and investigating
population genetics and evolution
of social animals. Methods of
genetic connectedness and family
structure provide significant and
new information regarding insects
‘breeding systems (Seppae &
Pamilo, 1995). For example,
cohabitation of many queens
(polygyny), male outputs by
workers, numerous matings by
females (polyandry), as well as
rapid queen turnover (Seppä et al.,
2009) are interesting phenomena.
In addition, the evaluation of
comprehensive genealogical
relationships among specified
individuals or populations of
individuals are the foundation to
test inclusively different fitness
forecasts (DeHeer & Herbers,
2004) and have provided some of
the most convincing evidence for
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 18
this (Pamilo, et al., 1997).
Population-level genetic
occurrences in ants are greatly
influenced by genetic structure and
the social nature of the ants ‘groups
that creates a direct connection
among the hierarchical levels. In
monogynous populations and
species (with one queen) and feebly
polygynous colonies, this is
projected to conclude in huge
homogenous groups and panmixia
in a huge area (Tsutsui & Case,
2001).
Conversely, in polygynous
species as well as populations,
mating takes place near or in the
nest. In addition, females are
frequently philopatric. This tends to
lead to a powerful population
structure, mainly in the
mitochondrial genome, as well as
raising the potential of regular
mating (Seppae & Pamilo, 1995). A
polygynous population form can
lead to a stable and patchy habitat
diminishing the success of
dispersing females (Pamilo et al.,
1997).
1.2.2 Migration
Migration is the transfer of species
from one place to another. Even
though it can take place in
recurring patterns (such as in
birds), migration when applied in a
population genetics context
regularly refers to the transfer of
species into or out of a distinct
population.
How does movement affect
relative allele frequencies (DeHeer
& Herbers, 2004)? If the moving
species stay and mate with the
indigenous individuals, they are
able to offer an unexpected influx
of alleles. Following the
establishment of mating among the
migrating and indigenous
individuals, gametes carrying
alleles are contributed by the
migrating individuals. This can
change the existing percentage of
alleles in the indigenous
population. Hence, movement out
of or into a population might be
accountable for a noticeable
alteration in allele frequencies (the
percentage of members having a
specific variant of a gene).
Immigration can also end in the
adding up of novel genetic variants
to the existing gene pool of a
specific population or species. In
addition, several factors impact the
amount of flow of genes among
diverse populations of ants. One of
the key features is movement, as
better movement of a species tends
to provide it better migratory
possibilities (Pamilo et al., 1997).
Sustained gene flow among
diverse populations can, in
addition, lead to a mixture of
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 19
different gene pools, dropping the
genetic difference of the two
cohorts. Because of this, gene flow
robustly represents in opposition to
speciation, via re-joining the gene
pools of the populations,
consequently, repairing the
devolving variances in genetic
differences that should have led to
complete speciation and formation
of a daughter species (DeHeer &
Herbers, 2004). For instance, if
types of ants inhabit one side of a
highway, ants may migrate from
one side to the other and vice versa.
If this ant is able to mate with the
other ant and generate viable
offspring, the alleles in the ant can
go from the group on one side of
the road to the other.
1.2.3 New colony development
One common feature shared
among ant colonies provides an
obvious criterion for colony
definition and supports the
possibility of a super-colony
staying as a single society: the
shared recognition cues of ants.
Studies of the Argentine ant
(Linepithema humile) display this
factor most successfully. The
Argentine ant lives in aggressive
super-colonies usually comprising
billions of queens and workers,
which can over to hundreds of
kilometers inside. A single colony
can spread through territories to
appropriate habitation due to a lack
of well-matched competitors. The
characteristic of super-colonies,
thus, is defined as the ability for
unlimited growth. Moreover, no
evidence has yet arisen to suggest
that the methods of the worker, the
indigenous patchiness of nests, and
the diet traffic in these extensive-
ranging populations are
consequently invariant that no
existing of super-colonies,
nonetheless as an alternative are
groups of several independent nest
collections that would be named as
(colonies) (Moffett, 2012).
1.2.4 Genetic structure
Due to physical barriers to
movement, by tending inadequately
for species to either alteration or
spread (vagility), as well as
propensity to stay or return to natal
habitat, normal populations hardly
entire breed as visualized, in
theoretical random models (Buston,
et al., 2007). A normal geographic
range is one in which species are
more intimately correlated to each
other compared to individuals that
are randomly chosen from the
whole population; this is illustrated
as the degree to which a group is
genetically structured. Genetic
structuring may be caused by
movement owing to species range
expansion, historical climate
change, or present availability of
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 20
habitat (Pamilo et al., 1997).
The structure of genetic
populations of species is affected by
internal (species- specific) and
external factors that can interrelate.
Internal factors contain inbreeding,
genetic drift, and dispersal
capabilities because of differences in
population bulks (Clemencet, et al.,
2005; Ruda, et al., 2010) Climate
and landscape affect the structure of
population in terms of the carrying,
fragmentation, and distribution
ability of appropriate habitats.
Christiansen and Reyer (2011) and
Zachos and Hartl (2011) have shown
how the structure of populations
may be controlled if gene flow is
decreased because of geographic
obstacles such as deserts, islands, or
due to disintegration in human-
inhabited landscapes. Additionally,
population structure may be affected
by historical phenomenon; for
example, ice ages or the range
spreading out from relict populations
(Grant, et al, 2011; Schmitt & Seitz,
2001).
Hölldobler and Wilson (1990)
found that distance, which happens
once the opportunities of mating
reduce among species with
geographical distance, affected how
distinctive the structure of a
population pattern was in
permanent habitats. The level of
this method relies on the spreading
abilities of the central organisms.
Additionally, in ants, winged
sexual scatter in mating flights of
distances from 50 meters up to a
few kilometers. For instance,
genetic investigates discovered
typical dispersion distances of 65 to
86 m as well as isolation by
distance (IBD) on this scale (Suni
& Gordon, 2010). In ants,
dispersion and mating through
courtship flights trailed via
autonomous colonies
‘establishment of winged queens is
evidently the ancestral reproductive
method (Foitzik, et al., 2011).
However, many ant species
have developed diverse mating and
dispersion behaviour even in
dependent colony foundations and
in nest mating through which no
wing queens escorted via workers
create new nests near the mothers
‘nest (Peeters & Ito, 2001).
Particularly, this substitutionally
generative style of colony fission or
budding, which is often revealed by
polygynous (Mult-queen) colonies,
leads to powerful organizing of ant
species on a local scale. It produces
a great relatedness between
neighboring nests as well as
diminishing relatedness upon the
remoteness of numerous meters.
This type of population stickiness
has been depicted in several ant
types with dependent colony
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 21
foundations (Seppae & Pamilo,
1995; Giraud, et al., 2000).
Restricted dispersal results in
an elevated relatedness between
mating partners. This, in turn, leads
to a high level of homozygosity
between progeny. This bears
suitability costs, for example, the
generation of sterile diploid males
in social Hymenoptera (J. M. Cook,
1993; Santomauro, et al., 2004),
decreased colony expansion and
survival, altered sexual investment
and smaller body size (Gerloff et
al., 2005; Tarpy & Page, 2002).
However, there has been a
detection of inbreeding and diploid
males in a number of ant species
with local dispersion (Haag-
Liautard, et al., 2009; Cole &
Wiernasz, 1997; Foitzik & Heinze,
2001; Sundström, et al., 2003).
Furthermore, in ants of the genera
hypo Ponera and Cardiocondyla
and in numerous invasive ant
species (Yamauchi, et al., 1996;
Yamauchi et al., 2001; S. Foitzik,
et al., 2002), normal intranidal (in-
nest) mating occurs (Foitzik et al.,
2011).
2. The meat ant (Iridomyrmex
purpureus)
The meat ant is the most
predominant Australian ant in
terms of its influence, abundance,
and effectiveness overcoming other
species of the surface fauna
(Greenslade, 1976). Due to the high
rates of their activity, their great
abundance, and their extreme
aggressiveness (Andersen & Patel,
1994) they are highly successful.
The meat ant is also known as the
gravel ant and it is also one of the
60 species of Iridomyrmex in
Australia (Shattuck & Barnett,
2001). It usually has a reddish head
and trunk contrasting with gaster,
and it is large (>4mm) (Andersen,
2000). There are at least eight color
forms of meat ants and together
they make up a very conspicuous
part of Australian ant fauna
(Halliday, 1983). Meat ants 'nests
are often large and built
underground with sand or gravel
mounded around the entrances to
the nest. The meat ants are often
involved in mutually beneficial
relationship with caterpillars. The
caterpillars provide sugary fluid for
the ants in return for protection
from predators (Shattuck, 1993).
The meat ant (Iridomyrmex
purpureus) is a dominant ant in
south eastern Australia (Andersen
& Patel, 1994), and it is one of the
60 species of Iridomyrmex in
Australia (Shattuck & Barnett,
2001). The meat ants ‘colony is
polygynous (a mating pattern in
which a single individual mates
with more than one individual of
the opposite sex) (Hölldobler &
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 22
Carlin, 1985), but less than 20% of
the galleries are controlled by
queens (Greaves & Hughes, 1974).
Hölldobler and Wilson (1990)
suggested, however, that many
mature nests consist of more than
one functional (egg-laying) queen
(Hölldobler & Carlin, 1985).
Workers (sterile females) perform
all other colony activity such as
feeding the young, serving the
queen, collecting water and food,
and protecting the nest (Wilson,
1976). Workers are also genetically
recognizable as solitary family
units combined to form huge multi-
nest colonies with a considerable
exchange of workers among the
nests (Halliday, 1983). Workers
perform most jobs within the
colony: some assist young queens
to establish colonies and others
attack strangers, indicating that
they might be able to discriminate
between outsiders and their nest
mates (Carew, et al., 1997).
In Australia, meat ants
(Iridomyrmex purpureus) are one
of the dominant ant species in ant
communities of eastern Australia
(Greenslade, 1976). Some of this
ecological success could be due to
their population genetics.
Population genetics provides
insight into the roles the essential
evolutionary forces of gene flow,
drift, and selection play in
processes such as speciation and
adaption (Ross, et al., 1999). Once
a population is isolated from
another, the accumulation of novel
mutations, genetic drift, and locally
differing selection can lead to
genetic variations between
populations. There are some factors
that may impact the altering of the
genetic structure of the meat ant I.
purpureus. Several theoretical and
empirical studies have
demonstrated that the population
structure of species is the result of
dispersal, mortality, and mating of
individuals (Rüppell, et al, 2003),
as well as the effects of breeding
(Storz, et al., 2001), local
extinction (Wade & McCauley,
1988), and restricted dispersal
(Hansson, et al, 2002). Assessing
population genetics does not only
investigate gene behaviour within a
population, it also includes the
study of how organisms adapt to a
stabilizing or fluctuating
environment (Cook, 1976).
2.1 Meat ant activity
The activity of a meat ant
infertile females is affected by
temperature so, in the summer,
workers search for food from dawn
to the late afternoon, while their
activity, in the winter, begins after
sunrise and continues until sunset
(Greenaway, 1981). Each day, the
large groups of infertile females
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 23
spend the day outside the nest that
in distance for feeding on
honeydew and excreting, while the
other rest of group spends the day
scavenging the territory around the
colony for proteinaceous and dead
insects (Greaves & Hughes, 1974).
The workers gather food from dead
animals by stripping off their meat
as the colonies of meat ants are not
far from each other (Mahboba,
2023). Nevertheless, their major
source of food is honeydew from
sap sucking insects (Greenslade &
Halliday, 1982). The meat ant's
food types depend on carbon or
nitrogen they have which getting by
colonies' surrounded soil
(Mahboba,2023).
2.2 Meat ants’ behaviour
and organization in colonies
Unlike other ant species, meat
ants do not have specific duties for
each ant; meat ants have various
roles in their colony during various
stages in their lives. This is known
as age cast polytheism. In their
colonies, it is the duty of the
younger ants to take care of eggs
and the larvae. Elderly ants make
up the bulky foraging groups for
exploiting substantial constant food
resources like dead animals. The
older ants search for their food
through open ground by gathering
invertebrates to be used in the
colonies (Shattuck, 1993). Meat
ants’ foraging and communication
with other insects Meat ants are
scavengers and predators. They are
diurnal;however, foraging becomes
bimodal during the hot season
(McIver, 1991). Like other species,
the meat ant engages in a symbiotic
relationship with caterpillars and
butterflies. The caterpillars and the
butterflies provide secretions that
are the ant ‘s food and in return, the
ant provides security from their
predators. Honeydew is the main
source of nourishment for the meat
ants. However, the honeydew can
be supplemented by the meat from
the dead animals (Schultheiss &
Nooten, 2013).
The meat ants can be very
aggressive towards ants from other
colonies (Anderson & Patel, 1994).
The meat ants engage the other ants
in battle to establish borders or
foraging barriers (Schultheiss &
Nooten, 2013). The meat ants are
so aggressive so when somethings
are closer from their colonies, they
will attack them (Mahboba,2023).
However, meat ants from larger
colonies are less aggressive to other
ants from small colonies
(Ettershank & Ettershank, 1982).
They can even recognize their mate
nest from other nests (Van
Wilgenburg, et al., 2006).
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 24
Furthermore ants ‘nests are built in
a homogenous environment, so this
environment may have similar food
resources (Mahboba, 2023).
2.3 Regulation of sexuality
Queen, workers (fertile,
infertile ants), and soldiers are
consisted the nest, and each has
different duties, such as the queen
is responsible for reproduce
(Hölldobler & Carlin, 1985), while
the works (infertile ) help the queen
during the reproductive process,
and also the queen is able to choose
and select the appropriate mate, and
the soldiers are mainly responsible
for protect the nest, specially, to
ensure all conditions are favorable
for the queen during mating
time(Keller & Wilson, 1993);
because of choosing the worker
who live in soil rich with (N)
(Mahboba,2023).
In addition, the queen
produces a hormone that helps to
identify and attract her mating
partner during the reproductive
season. Once the pheromone is
produced, it induces the worker
ants (infertile) into a reaction
(Mahboba,2023).
The hormone is also used to
help in the reproductive
development of the ants; it helps to
create the sexual ants. This helps to
keep colonies intact since the
mating is only limited to a few ants.
In some cases, it is believed that the
worker ant helps in determining the
sexual partner for the queen in
order to create a strong colony
(Crozier & Pamilo, 1996).
CONCLUSION
Meat ants, or Iridomyrmex
purpureus, are a prominent species
in ant populations located in the
eastern region of Australia. Some
of their ecological success might be
attributable to their population
genetics.
The study of population
genetics sheds light on the
functions that selection, drift, and
gene flow three fundamental
evolutionary forces play in
phenomena like speciation and
adaptation.
Genetic variances across
populations can result from the
accumulation of new mutations,
genetic drift, and regionally varying
selection when a population is
isolated from another.
ACKNOWLEDGEMENT
I would like to thank everyone help in
performing this work.
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 25
. REFERENCES
Andersen, A. N. (2000). The ants of northern
Australia: A guide to the monsoonal
fauna. East Melbourne: CSIRO.
Andersen, A. N. (2002). Common names for
Australian ants (Hymenoptera:
Formicidae). Australian Journal of
Entomology, 41(4), 285-293.
Bjostad, L. B., & Roelofs, W. L. (1986). Sex
pheromone biosynthesis in the red-
banded leafroller moth, studied by
mass-labeling with stable isotopes and
analysis with mass spectrometry.
Journal of Chemical Ecology, 12(2),
431-450.
Bourke, A. F. G., & Franks, N. R. (1995).
Social evolution in ants. Princeton, N.J:
Princeton University Press. Burwell, C.
J. (2007). Ants of Brisbane: Queensland
Museum.
Buston, P. M., Bogdanowicz, S. M., Wong, A.,
& Harrison, R. G. (2007). Are
clownfish groups composed of close
relatives? An analysis of microsatellite
DNA variation in Amphiprion percula.
Molecular Ecology, 16(17), 3671-3678.
doi: 10.1111/j.1365-294X.2007.03421.x
Chamberlain, P. M., Bull, I. D., Black, H. I.,
Ineson, P., & Evershed, R. P. (2004).
Lipid content and carbon assimilation in
Collembola: Implications for the use of
compound-specific carbon isotope
analysis in animal dietary studies.
Oecologia, 139(3), 325-335.
Charlesworth, B., Charlesworth, D., & Barton,
N. H. (2003). The effects of genetic and
geographic structure on neutral
variation. Annual Review of Ecology,
Evolution, and Systematics, 99-125.
Christiansen, D. G., & Reyer, H. U. (2011).
Effects of geographic distance, sea
barriers and habitat on the genetic
structure and diversity of all-hybrid
water frog populations. Heredity,
106(1), 25-36. doi:
10.1038/hdy.2010.37
Clemencet, J., Viginier, B., & Doums, C.
(2005). Hierarchical analysis of
population genetic structure in the
monogynous ant Cataglyphis cursor
using microsatellite and mitochondrial
DNA markers. Molecular Ecology,
14(12), 3735-3744. doi:
10.1111/j.1365-294X.2005.02706.x
Cole, B. J., & Wiernasz, D. C. (1997).
Inbreeding in a lek-mating ant species,
Pogonomyrmex occidentalis.
Behavioral Ecology and Sociobiology, 40(2),
79-86. doi: 10.1007/s002650050318
Cook, J. M. (1993). Sex determination in the
Hymenoptera: A review of models and
evidence. Heredity, 71(4), 421-435. doi:
10.1038/hdy.1993.157
Cook, L. M. (1976). Population Genetics:
Chapman and Hall. London.
Crozier, R. H., & Pamilo, P. (1996). Evolution
of social insect colonies. New York:
Oxford University.
Dalecky, A., Ponsard, S., Bailey, R. I.,
Pélissier, C., & Bourguet, D. (2006).
Resistance evolution to Bt crops:
Predispersal mating of European corn
borers. PLoS Biology, 4(6), e181.
DeHeer, C., & Herbers, J. (2004). Population
genetics of the socially polymorphic ant
Formica podzolica. Insectes Sociaux,
51(4), 309-316.
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 26
Detrain, C., Natan, C., & Deneubourg, J.-L.
(2001). The influence of the physical
environment on the self- organised
foraging patterns of ants.
Naturwissenschaften, 88(4), 171-174.
Ehleringer, J. R., & Cerling, T. E. (2002). C3
and C4 photosynthesis. Encyclopedia of
Global Environmental Change, The
Earth system: Biological and ecological
dimensions of global environmental
change, 2, 186-190.
Ettershank, G. (1968). The three-dimensional
gallery structure of the nest of the meat
ant Iridomyrmex Purpureus (SM.)
(Hymenoptera: Formicidae). Australian
Journal of Zoology, 16(4),715-723.
Evans, R. D. (2007). Soil nitrogen isotope
composition. Stable Isotopes in Ecology
and Environmental Science, 2, 83-98.
Fernandez, F., & Delsinne, T. (2013). Ant
ecology. Revista Colombiana De
Entomologia, 39(1), 174.
Fiedler, K., Kuhlmann, F., Schlick-Steiner, B.,
Steiner, F., & Gebauer, G. (2007).
Stable N-isotope signatures of central
European antsassessing positions in a
trophic gradient. Insectes Sociaux,
54(4), 393-402.
Fischer, K., O'Brien, D., & Boggs, C. (2004).
Allocation of larval and adult resources
to reproduction in a fruit‐feeding
butterfly. Functional Ecology, 18(5),
656-663.
Fischer, R. C., Richter, A., Wanek, W., &
Mayer, V. (2002). Plants feed ants: food
bodies of myrmecophytic Piper and
their significance for the interaction
with Pheidole bicornis ants. Oecologia,
133(2), 186- 192.
Fischer, R. C., Wanek, W., Richter, A., &
Mayer, V. (2003). Do ants feed plants?
A 15N labelling study of nitrogen
fluxes from ants to plants in the
mutualism of Pheidole and Piper.
Journal of Ecology, 91(1), 126-134.
Fisher, B. L., Sternberg, L. da S. L., & Price,
D. (1990). Variation in the use of orchid
extrafloral nectar by ants. Oecologia,
83(2), 263-266.
Foitzik, S., & Heinze, J. (2001).
Microgeographic genetic structure and
intraspecific parasitism in the ant
Leptothorax nylanderi. Ecological
Entomology, 26(5), 449-456.
Foitzik, S., Heinze, J., Oberstadt, B., &
Herbers, J. M. (2002). Mate guarding
and alternative reproductive tactics in
the ant Hypoponera opacior. Animal
Behaviour, 63(3), 597-597. doi:
10.1006/anbe.2001.1945
Foitzik, S., Rüger, M. H., Kureck, I. M., &
Metzler, D. (2011). Macro- and
microgeographic genetic structure in an
ant species with alternative reproductive
tactics in sexuals. Journal of
Evolutionary Biology, 24(12), 2721-
2730. doi: 10.1111/j.1420-
9101.2011.02397.x
Folgarait, P. J. (1998). Ant biodiversity and its
relationship to ecosystem functioning:
A review. Biodiversity and
Conservation, 7(9), 1221-1244.
doi:10.1023/A:1008891901953.
Frank, D. A., & Evans, R. D. (1997). Effects of
native grazers on grassland N cycling in
Yellowstone National Park. Ecology,
78(7), 2238-2248.
Gammans, N., Bullock, J. M., & Schönrogge,
K. (2005). Ant benefits in a seed
dispersal mutualism. Oecologia, 146(1),
43-49.
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 27
Gannes, L. Z., O'Brien, D. M., & del Rio, C.
M. (1997). Stable isotopes in animal
ecology: assumptions, caveats, and a
call for more laboratory experiments.
Ecology, 78(4), 1271-1276.
Gebauer, G., & Taylor, A. (1999). 15N natural
abundance in fruit bodies of different
functional groups of fungi in relation to
substrate utilization. New Phytologist,
142(1), 93-101.
Gerloff, C. U., & Schmid‐Hempel, P. (2005).
Inbreeding depression and family
variation in a social insect, Bombus
terrestris (Hymenoptera: Apidae).
Oikos, 111(1), 67-80. doi:
10.1111/j.0030- 1299.2005.13980.
Gibb, H., & Cunningham, S. A. (2011). Habitat
contrasts reveal a shift in the trophic
position of ant assemblages. Journal of
Animal Ecology, 80(1), 119-127.
Giraud, T., Blatrix, R., Poteaux, C., Solignac,
M., & Jaisson, P. (2000). Population
structure and mating biology of the
polygynous ponerine ant
Gnamptogenys striatula in Brazil.
Molecular Ecology, 9(11), 1835-1841.
doi: 10.1046/j.1365-294x.2000.01085.x
Grant, W. S., Merkouris, S. E., Kruse, G. H., &
Seeb, L. W. (2011). Low allozyme
heterozygosity in North Pacific and
Bering Sea populations of red king crab
(Paralithodes camtschaticus): Adaptive
specialization, population bottleneck, or
metapopulation structure? ICES Journal
of Marine Science: Journal du Conseil,
68(3), 499-506.
Gratton, C., & Forbes, A. E. (2006). Changes
in δ 13C stable isotopes in multiple
tissues of insect predators fed
isotopically distinct prey. Oecologia,
147(4), 615-624. doi: 10.1007/s00442-
005-0322-y.
Greaves, T. (1971). The distribution of the
three forms of the meat ant
Iridomyrmex purpureus (Hymenoptera:
Formicidae) in Australia. Journal of the
Australian Entomological Society, 10,
15-21.
Greaves, T., & Hughes, R. (1974). The
population biology of the meat ant.
Australian Journal of Entomology,
13(4), 329-351.
Greenaway, P. (1981). Temperature limits to
trailing activity in the Australian arid-
zone meat ant iridomyrmex pupureus
form viridiaeneus. Australian Journal of
Zoology, 29(4), 621-630.
Greenslade, P. (1974). Some relations of the
meat ant Iridomyrmex purpureus
(Hymenoptera: Formicidae) with soil in
South Australia. Soil Biology and
Biochemistry, 6(1), 7-14.
Greenslade, P. (1976). The meat ant
Iridomyrmex purpureus (Hymenoptera:
Formicidae) as a dominant member of
ant communities. Australian Journal of
Entomology, 15(2), 237-240.
Greenslade, P., & Halliday, R. (1982).
Distribution and speciation in meat ants
Iridomyrmex purpureus and related
species (Hymenoptera: Formicidae). In
W. R. Barker and P. J. M. Greenslade
(Eds.) Evolution of the flora and fauna
of arid Australia.
Haag-Liautard, C., Vitikainen, E., Keller, L., &
Sundström, L. (2009). Fitness and the
level of homozygosity in a social insect.
Journal of Evolutionary Biology, 22(1),
134-142. doi: 10.1111/j.1420-
9101.2008.01635.
Hahn, M., & Maschwitz, U. (1985). Foraging
strategies and recruitment behaviour in
the European harvester ant Messor
rufitarsis (F.). Oecologia, 68(1), 45-51.
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 28
Halaj, J., Peck, R. W., & Niwa, C. G. (2005).
Trophic structure of a macroarthropod
litter food web in managed coniferous
forest stands: A stable isotope analysis
with δ 15N and δ 13C.Pedobiologia -
International Journal of Soil Biology,
49(2), 109-118. doi:
10.1016/j.pedobi.2004.09.002.
Halliday, R. (1983). Social organization of
meat ants Iridomyrmex purpureus
analysed by gel electrophoresis of
enzymes. Insectes Sociaux, 30(1), 45-
56.
Halliday R.B., 1978. Genetic Studies of Meat
Ants (Iridomyrmex purpureus).
Unpublished Ph.D. thesis, University of
Adelaide.
Hansson, B., Bensch, S., Hasselquist, D., &
Nielsen, B. (2002). Restricted dispersal
in a long-distance migrant bird with
patchy distribution, the great reed
warbler. Oecologia, 130(4), 536-542.
Hershey, A. E., Pastor, J., Peterson, B. J., &
Kling, G. W. (1993). Stable isotopes
resolve the drift paradox for baetis
mayflies in an Arctic River. Ecology,
74(8), 2315-2325.
Hoering, T. (1955). Variations of Nitrogen-15
Abundance in Naturally Occurring
Substances. Science (New York, N.Y.),
122(3182), 1233-1234. doi:
10.1126/science.122.3182.1233
Högberg, P. (1997). 15N natural abundance in
soil‐plant systems. New Phytologist,
137(2), 179-203.
Hölldobler, B., & Carlin, N. F. (1985). Colony
founding, queen dominance and
oligogyny in the Australian meat ant
Iridomyrmex purpureus. Behavioral
Ecology and Sociobiology, 18(1), 45-
58.
Hölldobler, B., & Wilson, E. (1990). The Ants.
Cambridge, MA: Belknap: Harvard
Press.
Hood-Nowotny, R., Mayr, L., & Knols, B. G.
(2006). Use of carbon-13 as a
population marker for Anopheles
arabiensis in a sterile insect technique
(SIT) context. Malaria Journal, 5(1), 6.6
Hood‐Nowotny, R., & Knols, B. G. (2007).
Stable isotope methods in biological
and ecological studies of arthropods.
Entomologia Experimentalis et
Applicata, 124(1), 3-16.
Hunter, M. D. (2009). Trophic promiscuity,
intraguild predation and the problem of
omnivores. Agricultural and Forest
Entomology, 11(2), 125-131.
Jung, S., & Lee, S. (2012). Molecular
phylogeny of the plant bugs
(Heteroptera: Miridae) and the
evolution of feeding habits. Cladistics,
28(1), 50-79.
Keller, L., & Wilson, E. O. (1993). Queen
number and sociality in insects. Oxford,
England: Oxford University Press.
Kiyashko, S. I., Imbs, A. B., Narita, T.,
Svetashev, V. I., & Wada, E. (2004).
Fatty acid composition of aquatic insect
larvae Stictochironomus pictulus
(Diptera: Chironomidae): Evidence of
feeding upon methanotrophic bacteria.
Comparative Biochemistry and
Physiology Part B: Biochemistry and
Molecular Biology, 139(4), 705-711.
Kohl, D. H., Shearer, G. B., & Commoner, B.
(1971). Fertilizer nitrogen: contribution
to nitrate in surface water in a corn belt
watershed. Science, 174(4016), 1331-
1334.
Larson, P. P., & Larson, M. W. (1967). Ants
observed. London: Hale.
Derna University Journal of Medical Sciences, 2(2): 11-29, 2024
Mahboba, 2024 29
Lobry de Bruyn, L. (1999). Ants as
bioindicators of soil function in rural
environments. Agriculture, ecosystems
& environment, 74(1), 425-441.
Macneale, K. H., Peckarsky, B. L., & Likens,
G. E. (2004). Contradictory results from
different methods for measuring
direction of insect flight. Freshwater
Biology, 49(10), 1260-1268.
Macneale, K. H., Peckarsky, B. L., & Likens,
G. E. (2005). Stable isotopes identify
dispersal patterns of stonefly
populations living along stream
corridors. Freshwater Biology, 50(7),
1117-1130.
Mahboba, S., Aldareh (2023). Stable Isotope
Analysis of the Meat Ant (Iridomyrmex
purpureus). Derna University Journal of
Medical Sciences, 1(1): 1-8, June 2023.
Malainey, M. (2011). Stable Isotopes A
Consumer's Guide to Archaeological
Science (pp. 35-44): Springer New
York.
Malausa, T., Bethenod, M., Bontemps, A.,
Bourguet, D., Cornuet, J., & Ponsard, S.
(2005). Assortative mating in sympatric
host races of the european corn borer.
Science (New York, N.Y.), 308(5719),
258- 260. doi:10.1126/science.1107577.
Markow, T., Anwar, S., & Pfeiler, E. (2000).
Stable isotope ratios of carbon and
nitrogen in natural populations of
Drosophila species and their hosts.
Functional Ecology, 14(2), 261-266.
McIver, J. (1991). Dispersed central place
foraging in Australian meat ants.
Insectes Sociaux, 38(2), 129-137.
McNabb, D. M., Halaj, J., & Wise, D. H.
(2001). Inferring trophic positions of
generalist predators and their linkage to
the detrital food web in
agroecosystems: A stable isotope
analysis. Pedobiologia, 45(4), 289-297.
=====================

 Iridomyrmex Purpureus


      


        


      

   


       
       
        
 
        

     
       
      
     
 
      

I. Purpureus
      
    Iridomyrmex purpureus
30
ResearchGate has not been able to resolve any citations for this publication.
Article
Full-text available
Ants ‘ability to use a great diversity of food sources contributes to their evolutionary and ecological success. However, this broad dietary niche along with their sociality makes this group of species notoriously difficult to examine with regard to nutritional ecology. To overcome this difficulty, stable isotopes have been used to assess the trophic position of ants in the food web. Stable isotope technology has greatly contributed to our understanding of the food web ecology of social insects, and can determine the dietary history of organisms. Furthermore, this method has been utilized successfully to study nutrient fluxes enriched with carbon and nitrogen isotopes. The aim of this study is to show if there is a relationship between δ13C and δ15N of meat ant nests and comparing the ration with soil adjacent to the ant nests. This study was carried out on 62 nests of meat ants (Iridomyrmex purpureus) and 62 samples of soil located adjacent to the nests in Armidale in New South Wales, Australia. The first ants' nest was sampled at UNE, Armidale, and the following 61 nests were sampled in a northerly direction away from UNE on a wide road verge/ travelling stock route. There were no significant differences between the C and N stable isotopes collected from soil and ants' nests and distance that nests were located away from one another; and there were no significant differences between δ13C and δ15N and the size of the ant's nest (based on the N-S nest length). Finally, the δ15N levels of meat ants were not correlated with their surrounding soil. Future study of measuring δ13C and δ15N levels to investigate particular interactions among ants and surrounding soil as well as with other resources that ants may feed on will be necessary to observe ants in a wide range of environments and that it will increase our understanding of ants ‘food webs.
Article
Full-text available
The colonization cycle hypothesis states that stream ecosystems would become depleted of insects if flying adults did not compensate for drifting immatures. Using long-term drift and benthic abundance data, we show that a Baetis mayfly nymph population moves downstream during development in the Kuparuk River in arctic Alaska. Baetis relative benthic abundance decreased from early to late season in an upstream unfertilized river section, while simultaneously increasing in the downstream fertilized section. Baetis nymphs drifted significantly more in the upstream unfertilized section, compared to the downstream fertilized section where food was more abundant. Approximately one-third to one-half of the nymph population drifted at least 2.1 km downstream during the arctic summer. A stable isotope tracer experiment and mathematical models show that about one-third to one-half of the adult Baetis population flew 1.6-1.9 km upstream from where they emerged. These results provide a quantitative test of the colonization cycle for the dominant grazer/collector in the Kuparuk River. Quantifying the colonization cycle is essential to understanding stream ecosystem function because offspring of downstream insects are needed for nutrient cycling and carbon processing upstream. Since downstream drift and upstream flight are important components in recovery of streams from disturbances, our results provide a quantitative method for predicting recolonization rates from downstream, essential to estimating recovery.
Article
Full-text available
Trailing activity is restricted to the hours of daylight and is further limited by the temperature of the air surrounding the ant (T*a) and by the net radiant heat load. Activity in winter begins well after sunrise and continues until late afternoon averaging c. 7.4 h per day. Midsummer activity occupies two distinct periods, one commencing at dawn and the other in late afternoon; the two together total about 6.8 h per day. The upper and lower values of T*a, between which ants are active are 43.5°C and 14°C. Radiant heat input is dissipated from the ants by convective heat loss.
Book
Ants are one of the most important faunal groups in Australia and are widely used as bioindicators in land monitoring and assessment programs. The Ants of Northern Australia will help in the identification of the 1500 or more ant species occurring in monsoonal Australia, an area which encompasses most of the northern third of the continent. Until now, no book has described the northern Australian ant fauna below genus level. Such a treatment is required to support and promote the numerous ecological studies involving ants, especially in the context of their use as bioindicators. The Ants of Northern Australia features original analyses of genera at the species-group level, and so has relevance throughout Australia. It treats all major species that have been described, as well as numerous others that remain undescribed.
Article
Abstract Most insects do not have common names, and this is a significant barrier to public interest in them, and to their study by non-specialists. This holds for even highly familiar insect groups such as ants. Here, I propose common names for all major native Australian ant genera and species-groups, as well as for many of the most abundant and distinctive species. Sixty-two genera, 142 species-groups and 50 species are given names. The naming system closely follows taxonomic structure; typically a genus is given a general common name, under which species-group and species names are nested.
Article
A technique for mass-labeling was developed to study sex pheromone biosynthesis in the red-banded leafroller moth,Argyrotaenia velutinana. With this technique, the pheromone components and all fatty acyl groups in the pheromone gland were analyzed for incorporation of label in the same analytic ran with gas chromatography-mass spectrometry, using chemical ionization and selected ion monitoring (GC-SIM-CI-MS). Sex pheromone glands were incubated with fatty acids or triacylglycerols labeled with at least three deuterium atoms or carbon-13 atoms. The results of these incubations support an interpretation in which hexadecanoate is chain shortened to tetradecanoate, which is desaturated to produce (E)- and (Z)-11-tetradecenoate precursors for the sex pheromone components (E)- and (Z)-11-tetradecen-1-yl acetate. Labeled (E)- and (Z)-11-tetradecenoyl groups in synthetic triacylglycerols were not incorporated into the sex pheromone components, perhaps indicating that this lipid class is not a donor of the immediate fatty acyl precursors in sex pheromone biosynthesis.
Article
Several polymorphic allozyme loci (Amylase, Esterase-1, Esterase-3) have been used as genetic markers for the analysis of some aspects of social behaviour in two colour forms (actually sibling species) of meat ants,Iridomyrmex purpureus. A small amount of data suggests that haploid males are probably produced from eggs laid by the queen, and not by the workers. It appears that a substantial proportion of colonies of bothI. purpureus sens. strict, and the blue form of meat ants, may contain more than one queen. InI. purpureus sens. strict, colony organization is flexible, ranging from small, territorially distinct colonies in which the workers are recognizable genetically as single family units, to very large multi-nest colonies with considerabe exchange of workers between nests. This flexibility is seen as a response to the very variable types of habitat in which this form of meat ant occurs.
Article
We investigated the effects of native ungulates on grassland N cycling in Yellowstone National Park by examining natural 15N abundance (δ 15N) of soils and plants inside and outside long-term (32-36 yr) exclosures. Across six topographically diverse sites, grazers increased δ15N of soil (0-20 cm) by 0.27‰, which was substantial considering that values for ungrazed soil ranged 2.4‰ (2.4-4.8‰). The magnitude of grazer 15N enrichment was positively related (r2 = 0.70) to the intensity of herbivore activity during the study, indexed by the amount of dung (g/m2) deposited at the sites. We also found that soil δ15N of ungulate urine and dung patches was significantly higher than that of control areas. Grazers probably increased soil δ15N by promoting N loss from the soil via leaching, ammonia volatilization, and/or denitrification. Each of these processes results in the removal of 15N depleted products from the soil and, consequently, 15N enrichment of the remaining soil. In contrast to soil results, grazers reduced plant 15N by an average of 0.7‰, probably due to isotopically light, soil NO3- (compared to soil NH4+) constituting a more important N source for plants in grazed grassland relative to those in ungrazed grassland. These findings indicate that native grazers increased N loss from this north-temperate grassland as a result of accelerated losses on urine- and dung-affected microsites and, potentially, from elevated N loss throughout the grazed landscape due to grazers promoting N cycling. Furthermore, these results suggest that herbivores increase plant NO3- assimilation, which may positively affect primary productivity in this grazed ecosystem.