Carpenter Ants - Emporia State University

THE KANSAS SCHOOL NATURALIST
ISSN: 0022-877X
Published by EMPORIA STATE UNIVERSITY
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Cover Photo: A mature colony of carpenter ants has thousands of workers , all
infertile sisters.
John H. Klotz is an assistant
entomologist in the Department of
Entomology, University of California
at Riverside, Riverside, CA 9252l.
Laurel D. Hansen teaches at Spokane
Falls Community College, Spokane,
WA 99224 and co-authored the
Kansas
School
Naturalist
on
"Collection and Maintenance of
Ants" Vol. 41, No. 1. Byron L. Reid is
an entomologist and lives at 5311
Oakview, Shawnee, KS 66216.
Stephen A. Klotz provided all
artwork and is Chief, Section of
Infectious Diseases, Kansas City
Veterans Affairs Medical Center,
Kansas City, MO 64218.
CARPENTER ANTS
by John H Klotz, Laurel D. Hansen, Byron L. Reid and Stephen A. Klotz
Carpenter ants are some 01 the most
lascinating of all ants. Entomologists have
studied many aspects 01 their lile and
discovered important relationships which
carpenter ants have with other living
organisms including humans. The primary
role carpenter ants play in our North
American forest ecosystems is only
beginning to be appreciated.
at all, preferring to nest under rocks, in the
soil, or in living or dead non-woody plants.
The first North American ant ever to be
described scientifically was the black
carpenter ant. In 1773, Baron Carl DeGeer,
a student of the great Swedish botanist
Linnaeus,
described
Formica
pennsylvanica, now known as Campono/us
This ant is
pennsylvanicus (DeGeer).
common in the eastern half 01 the United
States (Figure 1), and though large and
conspicuous, it goes unnoticed because of
its nocturnal lifestyle. Nocturnal living
likely arose in response to competition
from other ants and predation by birds.
Birds, except for owls, are active in
daytime.
WHAT IS A CARPENTER ANT?
Carpenter ants belong to the large genus
01 ants, Campono/us, which is distributed
worldwide in tropical and temperate
regions. The common name, carpenter ant,
is derived from the prelerence of some of
the more notable Camponotus species to
excavate nests in wood. However, most
species of Camponotus are not "carpenters"
Figure I. Economically important species of carpenter ants and their North American geographic distribution.
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in the biological control of forest insects
and the recycling of organiC and inorganic
nutrients.
LIFE CYCLE
A winged ant begins the life cycle.
Carpenter ant colonies are formed after
reproductive adults (winged virgin queens
and winged males) emerge from their nest
of origin for mating flights, usually during
the first warm days of spring. After mating,
the male dies. Each inseminated queen
selects a nesting site, often in a small cavity
in a stump or log, or perhaps under bark in
a standing tree. She then breaks off her
wings and within a few days lays her first
eggs. In two to three weeks, the eggs hatch
into larvae that are fed by the queen , who
mobilizes food stored in the flight muscles
of her thorax and in the fat bodies of her
abdomen. The queen does not leave the
nest to hunt for food during this time .
Instead, she remains to protect, feed, and
raise the brood (Figure 3). At the end of the
larval developmental period, the larvae
pupate and emerge as minor workers,
numbering some 10 to 25 individuals .
Workers, which are all female, assume the
functions of foraging, nest excavating, and
brood rearing. In two years, a population of
workers ranging in size from small minors
(6 mm) to large majors (13 mm) will be
present. The size of a worker is not
genetically determined; rather it is
dependent on environmental factors such
as larval nutrition. Winged ad ults
(reproductives) are produced in six to 10
year-old colonies when populations exceed
2000 workers . Mature, or parent colonies,
establish satellite colonies
nearby
whenever there is a need for more territory,
resources, or a drier, warmer nesting site
for development of their larvae and pupae.
The queen, workers, and small larvae are
always present in the parent colony
whereas the satellite colonies contain
workers, larger larvae, and pupae. Except
during winter diapause , workers travel
between various satellites of the colony
that are connected by well-defined trails
(discussed later).
Figure 2. Camponotus modoc starts the recycling
process in a forest in Washington. This dead tree is
being cored out by a colony of carpenter ants.
Although carpenter ants are common
insects in woodland habitats and often go
unnoticed , th eir activities have far-reaching
effects on forest communities and humans.
Their social behaviors, such as group
foraging and food sharing, support large
colonies which by virtue of the queen
ants's longevity, make them permanent
residents of forests. Among the many
species of ants , the carpenter ants stand
out in forest ecosystems as the dominant
insect. Their predation on other insects
affects the population dynamics and
distribution of the species they prey upon ,
and their nesting habits initiate the
degradation process of tree cellulose
(Figure 2) to a form usable by other plants
and animals. Thus, in the ecological web of
forest life, carpenter ants playa critical role
4
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HOW CARPENTER ANTS
FIND THEIR WAY AROUND
Existing in an area with several
different colonies and avoiding
aggressive
encounters
requires
carpenter ants to be familiar with their
home range. Since they are primarily
nocturnal, they rely heavily on
physical cues and chemical trails for
orientation to and from the nest. Well­
maintained physical trails and trunk
lines of carpenter ants serve as
roadways through vegetation and
debris (Figure 4). These trails are
reminiscent of the wide, cleared trails
of leafcutting ants, common to Central
and South America.
In extreme
northern latitudes, carpenter ant trails
will often go underground following
natural hollows, such as those left by
decaying tree roots in the soil. These
tunnels are usually 1.5 to 3.0
centimeters in diameter and may be as
deep as one meter below the earth's
surface.
Figure 3. A newly mated queen carpenter ant with
her first brood.
COLONY SIZE
Populations of ant colonies can
reach tremendous numbers.
For
example, over 50,000 workers have
been found in colonies of Camponotus
modoc (cover), a western carpenter
ant. This is a relatively small social
group in comparison with another
carpenter ant found in the West ,
Camponotus vicinus, whose colonies
may number over 100,000 workers .
Part of the explanation for the vast
difference in populations of colonies is
the presence of multiple queens , a
condition called polygyny, that is
common in C. vicinus colonies. As
many as 41 functional queens have
been collected in a single C. vicinus
colony. Most species of carpenter ants
are monogynous (possessing only one
queen) and as a consequence, the
colonies are smaller and require years
to reach maturity.
5
Figure 4. Carpenter ants maintain large trunk lines.
One trunk line is shown below in a grassed area.
Figure 5. Carpenter ants forage at night on a oak tree using a "moon compass" for orientation.
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Chemical
trails
consist
of
hydrocarbons produced in the hindgut
of the ant and deposited on the trail
surface.
These hydrocarbons are
pheromones and are deposited by the
ant when the tip of her abdomen is
dragged on the substrate for short
distances as she moves along the trail.
Pheromones are odorous compounds
produced by the ant to convey
information.
In the case of trail
pheromones, the compounds guide
ants to locations outside of the nest.
Heavy deposits build up over time on
heavily traveled trails forming "trunk
trails" or main transportation arteries
guiding foragers
to
resources .
Resources include aphid colonies
where ants collect honeydew, a favored
food rich in sugars and sought by many
different ant species.
Trail pheromones also recruit
nestmates to newly discovered food
resources . Based on the relatively
large size of the carpenter ant's
olfactory lobes located in the brain,
the sense of smell is clearly important.
Smell serves the ants well in their
nighttime activities. However, the
individual forager eventually must
leave the trunk trail to search for
resources using both touch and sight,
and to accommodate this, other
orientation cues are used.
"Structural guideline orientation" is
one such important cue for foraging
carpenter ants . Unlike the chemicals
in odor tails, structural guidelines are
tactile stimuli in the form of edges,
grooves, or crestlines provided by tree
bark, vines, branches , or roots on the
forest floor. Carpenter ants follow
elaborate detours along branches or
sidewalks rather than go in straight
lines. An ant's movement is more
efficient on smooth, uncluttered
guidelines compared to movement
along trails on the ground where turf
and surface features impose numerous
obstacles to the ant's passage. The
benefit of these structural detours is
the shortening of overall trip time.
Structural guidelines are the
lowest level of cue found in investigations
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Figure 6. How carpenter ants orient in their environment. Left side: moon compass orientation as well as
landmarks may be used. Right side: sun compass orientation and edges, such as the house or telephone line
going into the house may be used. Pheromone hydrocarbons may be used as well, especially on well-travelled
roadways as shown by the path.
of carpenter ants' orientation system. If
placed in total darkness, ants are unable
to negotiate shortcuts by using visual
cues and resort to tactile orientation
along structural guidelines. If total
darkness is momentarily interrupted by
an overhead view of the forest canopy or
another visual cue, ants switch to
another orientation method called
landmark orientation.
Landmarks
ants as a directional cue (Figure 5). Felix
Santschi, a French entomologist,
demonstrated sun compass orientation
in desert ants in Africa in 1911. He
reversed the direction of the sun using
mirrors and showed clearly that desert
ants do orient by means of the sun. A
similar mirror experiment on moonlit
nights gives similar results with
carpenter ants . Foraging carpenter ants
reverse their direction in response to a
change in the apparent position of the
moon caused by the introduction of
mirrors.
Within the assemblage of orientation
cues for carpenter ants, there is a built-in
redundancy. Foraging ants actually rely
on more than one orientation cue; for
instance, a forager uses an odor trail as
well as a light source to orient. As a
consequence, the ants possess back-up
cues with which they can orient in the
absence of anyone particular cue. This
arrangement provides carpenter ants
with the ability to forage in the
woodlands in daylight and total darkness
(Figure 6).
include any visually conspicuous object
such as a tree or shrub. Landmarks are
memorized in detail and guide ants to
and from the nest.
Canopy orientation is one type of
landmark orientation that carpenter ants
use in temperate forests and under low
light conditions of the night sky. Since
carpenter ants nest within trees, the use
of leaf canopy landmarks as cues may be
an adaptation to increase the likelihood
of ants returning to the nest tree after
foraging.
Carpenter ants show a strong
response to light at night. This suggests
that the moon is also used by carpenter
7
Figure 7. Aphid colonies are tended by carpenter ants for honeydew.
FEEDING HABITS
from May through July but are less
attractive when offered in August or
September. Conversely, carpen ter ants
recruit slowly to simple sugar or honey
baits in the spring, but any carbohydrate
source is rapidly depleted from July
through the end of colony activity at the
time of approaching winter. Carbohydrates
are used as an energy source by adults
throughout the year, but the mass
provisioning in the fall , before the onset of
diapause, may contribute to overwintering
survival.
The redundancy in orientation cues is
fortuitous as it allows foraging under most
environmental conditions. To feed a colony
of over 100,000 ants is a formidable task.
From large, centrally located nests,
foraging ants will fan out along trails
leading to various destinations within the
forest habitat.
Carpenter ants are
voracious predators of arthropods, such as
flies, caterpillars,
beetles, harvestmen
(daddy long-legs), and spiders. Carpenter
ants also collect honeydew from aphids
and can often be observed tending them
(Figure 7). Aphids are small plant-sucking,
soft-bodied true bugs which excrete
copious quantities of honeydew, which is
rich in sugars. Many species of ants are
attracted to aphid-infested trees, shrubs,
and herbaceous plants. Husbandry of
aphids by ants is usually viewed as
detrimental to the host plants because the
aphid population usually grows under the
ants' . protection and aphids damage plant
tissue.
On the other hand, aphid
husbandry maybe beneficial since the ants
kill many plant eating insects that destroy
the host plants.
The black carpenter ant has a distinct
cycle of food preferences . During the
spring and early summer, when brood
production is high, the ants have a strong
preference for proteins, which are fed to
the developing larvae For example, freshly
diced mealworms are mobbed by workers
OPTIMIZING FEEDING
Foraging theory suggests that animals
conserve energy during foraging. One
prediction of foraging theory is that , as the
distance between the nest and a food
source increases, foragers will become
more selective in their diet. In economic
terms, the ants must maximize caloric
"revenue" to compensate for their
increased "expenditures" incurred by
foraging at greater distances from the nest.
If given a choice of different concentrations
of sucrose sugar on a feeding station, a
colony of Camponotus pennsy!uanicus will
preferentially
gather
the
higher
concentration as the distance travelled to
the feeding station increases. Therefore,
carpenter ants follow in practice the
foraging theory of maximizing energy gains
through selectively feeding among different
resources.
8
Figure 8. Carpenter ants on occasio n go to war over disputed territory. Here two colonies form a battle line
on a fal len log.
Another
notable
battle
was
observed in a hardwood forest in
Indiana between two colonies of
Camponotus pennsylvanicus. Upon
cessation of combat, thousands of
dead ants lay at the base of a tree in
which one carpenter ant colony was
nesting. This colony was attacked by a
migrating colony in search of a new
nesting location. The previous night's
carnage was frozen in time , fallen
soldiers locked in combat with their
were
disfigured,
foes.
Ants
dismembered, decapitated, often
disarticulated, and mandibles still
gripped the legs of dead adversaries.
Mortally wounded survivors could be
seen moving about with abdomens
severed or missing.
TERRITORIAL ANTS
GO TO WAR
Carpenter
ants
are
fiercely
territorial and battle with unrelated
members of their own species and with
other ants . The causes of these
territorial wars are unknown. We have
witnessed a number of such conflicts
between neighboring colonies of
carpenter ants . One large-scale conflict
of carpenter ants occurred in a dense
pine forest in Idaho. A battle line 90
meters long was drawn between two
neighboring colonies of Camponotus
modoc, and fighting occurred on the
ground and stumps . Part of the battle
line included a log 20 meters long lying
on the ground upon which combatants
were exposed on the crest of the log.
The war was waged for two entire days
and nights (Figure 8).
9
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AVOIDING WAR
WHY ACTIVE AT NIGHT?
Ant battles are momentous and clearly
exceptional behavior.
Under normal
conditions, carpenter ant colonies live side
by side, coexisting even at high densities.
For example, in a one acre plot in Indiana,
six unrelated colonies with their satellites
nested in 22 trees . These trees, and the
nest sites and foraging resources they
represented, were perfectly partitioned
among these colonies. In Florida, a one
acre tract of sandhill harbored 20 nests
representing nine coexisting unrelated
colonies and 11 satellite colonies.
To determine the relatedness and
territory size of colonies, one relies on the
aggressive behavior of carpenter ant
workers toward non-nestmates . Aggression
between non-nestmate workers ranges
from mild encounters where the ants fence
with their mandibles (back cover) to
intense interactions where prolonged
combat and death result. In order to test
colony relatedness, worker ants from one
tree are paired in a plastic tube with
workers from a different tree (Figure 9). If
there is no aggression between the ants,
they are most certainly colony nestmates.
If there is mild or intense aggression, the
ants . are non-nestmates from different
unrelated colonies. These interactions
between ants from different trees or nests
can be used to draw accurate maps of the
individual colony distributions, including
satellite colonies.
By working after sunset (nocturnal),
carpenter ants are able to share resources
with competing species of ants that work
during the day (diurnal). For example, in
Kansas the
black carpenter ant,
Campana/us pennsylvanicus, and a species
of black field ant, Formica subsericea, live
together in woodland habitats and forage
for similar foods. During the daytime, the
field ant co llects aphid "honeydew" in the
same trees which are used later in the
evening by carpenter ants for the same
purpose .
Since carpenter ants are
prima rily nocturnal , and the field ant is
diurnal, a clear diviSion of daily rhythms
occurs. Moreover, for unknown reasons ,
carpenter ants are able to sustain a higher
travelling velocity than the field ant as
temperatures drop, another adaptation to a
nocturnal existence.
An additional advantage carpenter ants
gain from a nocturnal existence is avoiding
predators that rely on sight for spotting
their prey. The few carpenter ant workers
foraging during the day make large,
conspicuous prey for birds such as robins,
grackles, and starlings. Experimental set­
ups placed in the field to study the ants
must be designed to prevent birds from
feeding on the exposed carpenter ants.
Figure 9. An aggreSSive encounter between two carpenter ants from different colonies.
10
ECOLOGICAL VALUE
OF CARPENTER ANTS
Foraging carpenter ants are a
dominant force and vital link in the
forest food web.
The impact of
carpenter ants on a forest ecosystem is
immense. They play a key role by
serving as the premier natural
biocontrol agents of such forest
defoliators as tent caterpillars and
spruce budworms. However, since
most species of carpenter ants are
nocturnal, studies of predation are
difficult to conduct and are therefore
few in number.
Carpenter ants can also be
considered an indicator species of the
health of a forest.
For example,
pileated woodpeckers, Hylatomus
pileatus, are rarely or never seen in
forests without Camponotus modoc.
This carpenter ant selects nest sites in
decaying
stumps
and
current
monoculture techniques in our large
forests do not allow for a variety of
habitats such as decaying stumps.
Thus, pileated woodpeckers are not
seen in such managed forests. In the
western United States, the pileated
woodpecker, a crow-sized bird, will not
survive the winter without access to
trees, snags, or tree stumps containing
colonies of the carpenter ant since
carpenter ants constitute the bulk of
the winter diet of these large
woodpeckers (Figure 10).
Figure 10. The pileated woodpecker depends upon
the presence of carpenter ants as a protein source.
These pileated woodpecker chicks await the arrival of
one of their parents.
Figure I I. Many species of passerine or perching
birds occasionally "ant," often after new feather
growth. Here is shown a bluejay (Cyanocitta cristata)
engaging in passive anting with carpenter ants. Even
poultry, such as the peafowl may engage in anting. In
active anting, the birds place ants in their feathers with
their beaks.
On the other hand many species of
birds undergo "anting," a process
where ants (including carpenter ants)
are used for their excretions in the
bird's act of preening. The purpose of
this behavior is unknown but some
scientists speculate that it may serve
an anti-ectoparasitic or antibiotic
function (Figure 11).
11 SURVIVING WINTER
I
What happens to the ants during the
cold winter months ? Parent colonies
containing the queen, workers, winged
reproductives, and larvae overwinter in a
metabolic state termed diapause . In
temperate regions, diapause is a period of
dormancy during which the ants are in a
state of "suspended animation ." The
encasing wood of the colony's residence
provides the overwintering colony with
insulation from cold temperatures .
Carpenter ants also produce glycerol, a
compound which acts as an antifreeze
preventing destructive ice crystals from
forming in their bodies.
In temperate regions , colonies break
diapause from January to June (depending
upon the latitude, elevation, and habitat),
and the queen begins her first egg-laying
period of the season, lasting for 7-10 days.
The voracious appetites of the developing
larvae trigger increased foraging activity.
The most intense foraging of the season
occ urs when the workers are driven by
increasing food requirements of the rapidly
developing larvae. A second peak of
activity occurs in June when the queen
again lays eggs for a period of 7-10 days.
The foraging activity period in the second
peak is shorter and less intense, and the
colony enters into diapause in September
or October along with the late summer
brood, which overwinters as larvae and
completes development in February.
Colonies are perennial and may exist for
more than 20 years.
DESTROYING WOOD
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Species of Camponotus that live in forest
environments and serve as important
ecological components are also recognized
as structural or nuisance pests in human
habitations (Figure 12).
As is the case with many organisms,
human activities have greatly influenced
the distribution and
abundance of
carpenter ants. In the northern United
States and the provinces of Canada,
carpenter ants cause millions of dollars of
damage to structures and to standing trees
used for lumber. For instance, a minimum
of 50,000 houses are treated each year by
professional exterminators in the state of
Washington for carpenter ants, and many
12
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more are treated by homeowners
themselves. One example from Washington
will illustrate the damage that can be done
by Camponotus modoc. When an older
home was being remodelled, the inner
wallboards were completely removed . Most
of the wall studs along a 20-foot wall were
tunneled by carpenter ants. The most
seriously damaged wood was so
extensively tunneled that an 8-foot-Iong
two-by-four weighed less than two pounds.
This tunnelling also extended into the attic
joists so that the owner fell with one leg
through the ceiling while he was showing
the damage. A home near Grand Rapids ,
Minnesota, had sawdust piles eight to ten
inches in height from carpenter ant
excavations in the basement and attic.
All across suburban America, it is a
common practice to build houses on
forested lots without removing the trees.
Unfortunately, nearly all forested lots
contain one or more carpenter ant
colonies, and the newly constructed house
is frequently invaded by satellite colonies
even before construction is completed.
Thus, homeowners are not dealing with
colonies that grow from the progeny of a
Single queen, but their houses suffer from
the damage of 5,000 to 50,000 workers in
satellite colonies that move into the
structure. Since these houses and other
man-made structures have optimal
temperature and moisture conditions for
the rearing of brood , they are ideal satellite
nesting sites.
Notwithstanding
the
potential
destructiveness of carpenter ants, their
household invasion is sometimes a blessing
in disguise, at least from the point of view
of the scientist. One of the authors took
advantage of a carpenter ant infestation to
study ant behavior. The colony, whose nest
was outside in a tree, followed a telephone
wire into the house, through an electrical
switch plate and along the edge of a kitchen
counter to a loosely capped jar of honey. In
place of the honey, a mixture of sugar in
milk and diced mealworms were offered to
the foraging ants. After several days of
feeding on this rich resource, the a nt
colony migrated into the house and located
its new nest site in some moist wood next
CONTRAST BETWEEN
TERMITES AND
CARPENTER ANTS
Termites
Straight Antennae
Wings of Equal Size
Thick Waist
Elbowed Antennae
Wings of Unequal Size
Thin Waist
14
to a dripping pipe underneath the sink.
It was in this makeshift laboratory
setting that carpenter ants revealed
some of their special adaptations
which make them so successful not
only in their natural surroundings but
in our home environments as well.
Overall
the benefits from the
"premier forest predator" far outweigh
the harm caused when humans and the
carpenter ant meet on common
ground. Future studies will surely
uncover further secrets of carpenter
ant behavior and biology as well as
better management techniques. Living
together with carpenter ants will
continue to challenge both scientists
and homeowners .
REFERENCES:
Bull, E. L., R. C. Beckwith and R. S.
Holthausen. 1992. Arthropod Diet of
Pileated
Woodpeckers
in
Northeastern Oregon. Northwest
Naturalist 73: 42-45.
Ehrlich, P.R., D. S. Dobkin and D.
Wheye.
1986. The Adaptive
Significance of Anting. Auk 103: 835.
Hansen, L. D. and R. D. Akre. 1990.
Biology of Carpenter Ants . Pages
274-280 in R. Vander Meer, K. Jaffe
and A. Cedeno [eds.], Applied
Myrmecology: A World Perspective.
Westview, Boulder, CO.
Holldobler, B. and E. O. Wilson. 1990.
The Ants. Belknap, Cambridge, MA.
Holldobler, B. and E. O. Wilson. 1994.
Journey to the
Ants.
Belknap,
Cambridge, MA.
Klotz, J. H. and B. L. Reid. 1992. The
Use of Spatial Cues for Structural
Guidelines in
Tapinoma sessile
and Camponotus pennsylvanicus.
Journal of Insect Behavior 5: 71-82.
Klotz, J. H. and B. L. Reid. 1993.
Nocturnal Orientation in the Black
Carpenter
Ant
Camponotus
(DeGeer)
pennsylvanicus
(Hymenoptera: Formicidae). Insectes
Sociale 40: 95-106.
15
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KANSAS SCHOOL NATURALIST, Box 4050
EMPORIA STATE UNIVERSITY
1200 Commercial Street
Em poria, KS 66801-5087