COLONIZATION

Red harvester ant nests are characterized by a lack of plant

growth and small pebbles surrounding the entrance to the tunnel,
which usually descends at a pronounced angle. Hulls of seeds
may
however be found scattered around the nest.
In grassland areas, such as ranches, the lack of plant life makes
red harvester ant colonies very easy to spot, and where they are
very plentiful they may make serious inroads into the grazing
available to livestock.
The mounds are typically flat and broad, 0 to 100 mm (0.0 to
3.9 in) high, and 300 to 1,200 mm (12 to 47 in) in diameter. There
have been reports of even larger denuded areas, on the order of
10 m2 (110 sq ft). Three to eight trails typically lead away from
the mound, like "arms". These trails are used by ants to collect
and bring food back to the mound. "Scout" ants are the first ones
out of the mound every morning. They seek food, and mark their
path as they return to the mound to alert the worker ants. The
worker ants follow the scent trail and collect the food. Other
worker ants clean, extend and generally tend to the mound, the
queen and the brood. All the ants in the colonies are females
apart from the winged males produced in the breeding season.
Initial colony formation
The reproductive unit of ant populations is the colony. A single
virgin queen first mates with several males at a reproductive
aggregation site formed by male harvester ants. She then flies to
a new site to produce an offspring colony.
DIET
The main food source for red harvester ants usually consists of
seeds, which they hoard in great numbers. The food is first ground
to a bread-like consistency using the ant's large mandible, and is
then stored in a granary, assuring the colony access to food year-

round. Seed collection on behalf of the red harvester ants benefits

their ecosystem through the process of myrmecochory, in which
ants aid in the dispersal of seeds while foraging for food. Both
plant and ant benefit from this relationship: the plants increase
their dispersal range and density, while the ants benefit from
acquiring nutrients and ensuring a more secure food supply in
future harvests. This is typically understood as amutualistic
interaction. Dead insects are also collected during foraging.
FORAGING (mencari makan)
Much research has been done on the foraging behavior of the red
harvester ant. Three types of workers are most involved in the
foraging process: nest patrollers, trail patrollers, and foragers. On
a given day, nest patrollers emerge first from the nest to assess
the safety and profitability of foraging. The colony gets the
majority of its water from the metabolism of the fats in seeds.[4] If
food is scarce, or if it is a particularly hot day, the energy and
water benefits of foraging may be outweighed by its energy and
water costs. In this case, the colony may be forced to rely on its
extensive food stores. Seeds may be stored in the nest for months
or even several years.[5]
The
colony
is
able
to
communicate
through
momentary antennal contact
involving
the
transfer
of cuticular hydrocarbons.[6] Other
visual,temporal,
[6]
and olfactory cues may also be involved. Based on the nest
patrollers reports, trail patrollers may leave the nest to determine
the best possible foraging direction. This decision is based upon
various economic factors such as food availability and
neighboring nests foraging behavior. As both types of patrollers
return, foragers assess their rates of return to decide whether to
leave the nest to find food. In an experiment involving patroller
mimics, it was found that a return rate of 1 patroller every 10
seconds stimulated the highest level of foraging activity. [6] This
return rate indicates high availability of food and good foraging

conditions, and therefore a favorable cost-benefit ratio for the

foragers. If the patroller return rate is too high, it may be a
warning of danger, such as the sighting of a predatory lizard. A
lower return rate could indicate lack of available food, or heavy
competition.
Ant foraging is guided by chemical signals that lead the ants up to
5060 km (?) from the nest at times. [7][better source needed] Once an ant
has decided to forage, it will almost always continue until it has
found food to take back to the nest. 90% of returning foragers are
successful.[8] Therefore, forager return rate is a good indicator of
food availability. Logically, overall foraging activity is influenced
by the rate of returning foragers. [8] When food is plentiful, foragers
will quickly find it and immediately return to the nest. Foragers
still in the nest will interpret this to mean food searches will be
profitable: low energy input with a high chance of a seed reward.
Foraging activity will consequently be increased. Likewise, a
decline in food availability, indicated by a decrease in forager
return rate, will cause the colony to decrease its foraging activity.
[6]

The ability for a colony to quickly regulate its foraging behavior is

quite important since their food source is variable and scattered.
This regulatory ability is variable in itself, influenced by food
availability, current need to eat, and colony health. [2] Colonies are
more likely to adapt to varying forager return rates when the rate
of foraging is high, which may be because variances are simply
harder to detect during low rates. [2] Overall, the red harvester ant
demonstrates a remarkable ability in social cognition, cost-benefit
analysis, and behavioral economics.
Collective regulation[edit]
Harvester ant populations operate without a centralized control
system by means of a decentralized chemical communication
system, through the process of chemical signalling. Foraging is

regulated using positive and negative feedback loops. When ants

return to the colony, this stimulates outgoing foraging. When ants
return to the colony at a higher rate with seeds, this in turn
causes the outgoing rate of foragers to increase. As fewer ants
bearing seeds return to the colony, the rate of outgoing foragers
decreases. The result is foraging regulation relative to the
abundance of available food at any given time. [9] The interactions
between returning and outgoing foragers operates on a timescale
of seconds. It typically only took three to eight seconds for an
outgoing forager to leave the nest as a result of an increase in
successful returning foragers.[10]
Relative abundance of food available is gauged by the average
time spent foraging per seed, rather than the distance traveled to
find each seed. Foragers take less time when food is more
abundant. Faster foraging signals faster rates of outgoing
foragers, maximizing the harvest of food relative to the
abundance in the area. Seeds are used as a source of water for
ants, and are often kept in the nest for anywhere from a few
months to multiple years.[11]
Interference competition in foraging[edit]
Interspecies
competition exists
between
harvester
ant
populations, presenting challenges to successful foraging. One
example of this is the competition for access to seed resources
between
the Novomessor
cockerelli and Pogonomyrmex
barbatus species of harvester ants. Novomessor cockerelli ants
engage in interference competition againstPogonomyrmex
barbatus colonies by "plugging" entrances to the nest with sand
and small rocks. This delays activity of P. barbatus for anywhere
from one to three hours, shifting the onset of foraging until later
in the day when the temperature is substantially higher. This
reduces productivity in two ways; firstly, the ants are delayed and
consequently have less time to forage; secondly, high soil
temperatures prematurely drive harvest ants back into their

nests, shortening the range of possible foraging. Research

indicated that the affected Pogonomyrmex barbatus populations
did not take efforts to compensate for the interference in their
activities, such as by inducing higher rates of foraging. As such,
nest plugging by rival species impedes the overall productivity of
worker ants in the species Pogonomyrmex barbatus.[12]
Division of labor[edit]
Caste system[edit]
Red harvester ants are eusocial insects, characterized by a highlevel of cooperation and an organized division of labor. The worker
caste is sterile. As with most ant species, reproduction is
performed
by
a
mating
caste
that
consists
of
winged alates (reproductives) that reside in the nest until weather
permits them to fly away and mate. After copulationthe male
usually dies, while the now-fertilized queen returns to the ground
to search for a suitable nesting site. Once she has chosen a site,
she sheds her wings and begins to reproduce, creating a new
colony. She produces "worker ants" for 120 years until her death.
Some Queens have been known to live up to 30 years in the wild.
In Pogonomyrmex barbatus, reproductive and non-reproductive
tasks are divided into a caste system, as discussed above. Some
individuals are specialized for reproductive function while other
ants are responsible for non-reproductive roles such as defense,
caring for the brood, and foraging for food. Research indicates
that the system of division of labor naturally arises in conjunction
with the formation of the group, rather than as a secondary
adaptation; this was demonstrated by an experiment in which
otherwise solitary queen harvest ants were placed in forced
association. The results of the study indicated that divisions of
labor arose in groups that lack an evolutionary history of such
social arrangements.[13]
Non-reproductive caste[edit]

Within non-reproductive Pogonomyrmex barbatus worker ants,

five major tasks are performed: foraging, maintenance of the
nest, defense, midden work, and convening. Roles are not
constant and chemical interactions determine when certain ants
engage in certain activities. As such, these tasks are relatively
fluid and cannot be understood as separate processes. The
transition in behavior on behalf of one group triggers behavioral
transitions in other groups of worker ants. [14]
Reproductive caste[edit]
The reproductive Pogonomyrmex barbatus caste are responsible
for the formation of new colonies. A single queen ant establishes
every colony. Large numbers of winged male ants and virgin
queen ants fly to mating aggregation sites following mid-summer
monsoon
rains.
In
order
to
found
a
successful
colony, Pogonomyrmex barbatus queens must mate with males
from two separate lineages. One lineage results in ants that will
become the workers of the colony. These are the ants responsible
for the non-reproductive tasks of the colony, including: foraging
for food, building and defending the nest, and brood care. The
other lineage produces reproductive female ants. After mating
with the male ants at the aggregation site, the queen flies away
to establish a new colony. Upon arriving at her new nest site, the
queen loses her wings and builds a nest in the soil. The entrance
into the nest is then sealed off in order to lay her eggs in safety.
With this, the colony is born. Over the course of the following six
to eight weeks, the primary brood of non-reproductive worker
ants emerges and begins to forage and build the colony. [15]
Reproduction[edit]
Mating behavior[edit]
Mating is perennial and species-specific between sub-species of
the Red Harvester Ant. Males distinguish reproductive females of
their species from females of other species using pheromones.

This occurs as a result of sexually stimulating pheromones

secreted from the poison-glands of reproductive Pogonomyrmex
barbatus females. These pheromones are perfectly detected by
males of the species upon physical contact, resulting in the
stimulation of sexual behavior.
Mating occurs at reproductive aggregation sites. Secretions from
the mandibular glands of males initially mark these sites. The
secretions attract females, as well as additional males, resulting
in strong male competition for mates. The system of mating
within the Pogonomyrmex barbatus species is comparable
to lek behavior performed by manyvertebrate species. The male
ants congregate and collectively give off a pheromone that
attracts reproductive females. The more males present to give off
the pheromone, the stronger the attraction for the females.
Although this practice strongly favors certain males and leaves
many others without mates, the congregation of males attracts
more females on the whole as opposed to pheromone release on
behalf of a solitary male. Consequently, it is more desirable for
the less attractive males to remain in the lek than to attempt to
attract females on their own.[16]
Sexual selection[edit]
Within
reproductive
aggregations
of Pogonomyrmex
barbatus ants, strong male competition exists. Male harvester
ants engage in communal mating displays in order to attract
females. Male ants will attempt to mate with any female they
encounter, but females resist copulation from the males.
Consequently, the communal mating displays heavily favor
reproduction with larger or more persistent males who are able to
overpower the females and successfully copulate. The male ants
will copulate for as long as possible, likely to restrict access to the
female from other males to ensure their sperm survives to
reproduce.[17]

Predation[edit]
Red harvester ants are a favorite food for desert horned lizards.
Harvester ants will defend their colonies vigorously against real or
perceived attacks, whether by large or small animals. They may
bite ferociously and their stings are venomous and painful. The
effect spreads through the lymphatic system, sometimes causing
dangerous reactions, especially in animals sensitive or allergic to
their venom.
Over the years, their numbers have been declining, and this has
often been attributed to competition for food with the invasive red
imported fire ant and the Argentine ant. Their decline has affected
many native species, especially those for which the red harvester
ant is a chief source of food, such as the Texas horned lizard.