''Polyergus breviceps'' is a
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of
ant
Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
endemic
Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to the
United States
The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. It is a
social parasite of other ants, namely of ''
Formica gnava'' but also of ''
Formica occulta'' and ''
Formica argentea''.
''Polyergus'' is an
inquiline
In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms, such as insects, may live in the ...
parasite
Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
, having lost its ability to take care of its young and themselves. "The workers do not forage for food, feed the young or the queen, or even clean up their own nest".
To survive, ''Polyergus'' workers raid ''Formica'' nests to steal the
pupa
A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
e—which, once hatched, become workers of the mixed nest. This sort of relationship is not unique, of the approximately 8,800 species of ants, at least 200 have evolved some form of
symbiotic
Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biolo ...
relationship with one another.
What makes ''Polyergus'' special is the way a newly mated queen can, all by herself, take over a ''Formica'' nest and start a new colony.
Slave raids and colony takeovers
''Polyergus'' workers emerge from their
nest
A nest is a structure built for certain animals to hold Egg (biology), eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of ...
(a mixed nest where ''
Formica
''Formica'' is a genus of ants of the subfamily Formicinae, including species commonly known as wood ants, mound ants, thatching ants, and field ants. ''Formica'' is the type genus of the Formicidae, and of the subfamily Formicinae. The type ...
'' workers are already enslaved) and
forage
Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also used m ...
for a suitable raid target. If they find one, the
scouts
Scouting or the Scout Movement is a youth social movement, movement which became popularly established in the first decade of the twentieth century. It follows the Scout method of informal education with an emphasis on practical outdoor activi ...
return to the nest, rally the other ''Polyergus'', and head out in a raiding column. If the new target is a ''Formica'' nest, the ''Polyergus'' incite a "panic-flee" response from the ''Formica'' by releasing
formic acid
Formic acid (), systematically named methanoic acid, is the simplest carboxylic acid. It has the chemical formula HCOOH and structure . This acid is an important intermediate in chemical synthesis and occurs naturally, most notably in some an ...
and take the ''Formica''
pupae
A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...
back to their nest as
slave
Slavery is the ownership of a person as property, especially in regards to their labour. Slavery typically involves compulsory work, with the slave's location of work and residence dictated by the party that holds them in bondage. Enslavemen ...
s. Occasionally, a new queen goes on a raiding column with the other ''Polyergus''. She then, after mating, leaves the column and forages for a suitable ''Formica''
nest
A nest is a structure built for certain animals to hold Egg (biology), eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of ...
.
Howard Topoff evaluated how the queen takes over a colony. After finding a ''Formica'' nest, she finds an entrance and is immediately attacked by ''Formica'' workers. The queen responds by biting with her sharp
mandible
In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla).
The jawbone i ...
s and releasing a
pheromone
A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
from her enlarged
Dufour's gland
Dufour's gland is an abdominal gland of certain insects, part of the anatomy of the ovipositor or sting apparatus in female members of Apocrita. The diversification of Hymenoptera took place in the Cretaceous and the gland may have developed a ...
that, unlike many other parasitic ants, has a pacifying effect. The queen quickly searches for and locates the ''Formica'' queen and, with her adapted mandibles, proceeds to bite and lick various parts of the ''Formica'' queen for an average of 25 minutes.
"Within seconds of the host queen's death, the nest undergoes a most remarkable transformation".
The ''Formica'' workers cease to be aggressive to the intruder and start to groom the ''Polyergus'' queen as if it were their own.
The takeover now complete, the ''Polyergus'' queen gains not only a nest, but a worker
caste
A caste is a Essentialism, fixed social group into which an individual is born within a particular system of social stratification: a caste system. Within such a system, individuals are expected to marry exclusively within the same caste (en ...
as well. She then lays her eggs and the cycle continues.
Mechanisms
Nearly all slave-making ants, including ''Polyergus'', have mandibular adaptations that help them attack others.
Specific to ''Polyergus'', when the queen first enters a ''Formica'' nest she releases a pheromone from her enlarged Dufour’s gland. Topoff did experiments to show that this pheromone has an important facilitative effect in colony usurpation; it reduces the aggression of the defending ''Formica'' workers. The researchers took the Dufour’s, pygidial, and poison glands from freshly mated ''Polyergus'' queens, using water as a
negative control. These were crushed in distilled water to make a solution containing their extracts. Because it was impossible to prevent a ''Polyergus'' queen from secreting her own pheromones during a live encounter, they used the harvester ant ''
Pogonomyrmex occidentalis
''Pogonomyrmex occidentalis'', or the western harvester ant, is a species of ant that inhabits the deserts and arid grasslands of the American West at or below . Like other harvester ants in the genus '' Pogonomyrmex'', it is so called because o ...
'' (naturally attacked by ''Formica'') as the subject. These ants were dipped in the above solutions, one ant/solution/test at a time, and placed in a
petri dish
A Petri dish (alternatively known as a Petri plate or cell-culture dish) is a shallow transparent lidded dish that biologists use to hold growth medium in which cells can be cultured,R. C. Dubey (2014): ''A Textbook Of Biotechnology For Class- ...
with three ''Formica occulta'' "attackers". They were observed for three minutes to see for how long the ''Pogonomyrmex'' was attacked.
The results were as follows: "the mean duration of aggression by the Dufour’s gland treatment was 53.3 seconds...The mean duration of aggression for the water, pygidial gland, and poison gland controls were: 143.5 seconds, and 137.2 seconds and 132.2 seconds respectively".
Apparently, at some time, ''Polyergus'' queens evolved the capacity to passively facilitate colony takeovers.
Topoff and Ellen Zimmerli also did experiments to prove that the ''Polyergus'' queen "tricks" the ''Formica'' colony by obtaining chemicals from the ''Formica'' queen in the process of killing her. In one test, the ''Formica'' queen was killed (by flash freezing then thawing) prior to ''Polyergus'' contact. Even though it was already dead, the ''Polyergus'' queen bit, stabbed, and licked the queen just as if it were alive: and the ''Polyergus'' was consequently accepted by the colony.
Another test showed that if no ''Formica'' queen was present, then ''Polyergus'' had little chance of a successful takeover. Clearly, the ''Formica'' queen is providing some sort of chemical(s) to the ''Polyergus'' queen, however unintentionally. That takeover can occur even if the host queen is dead, but not if she is not present, proves that chemical absorption is important. The ''Polyergus'' queen needs only kill one host queen to be accepted. If the colony was
polygynistic, the ''Polyergus'' queen can take her time finding the other queens. "Hour by hour, day by day, she methodically locates and kills every ''Formica'' queen, sometimes taking several weeks to clear out all remnants of opposition".
Evolution
Considerable work has been published on the
evolution
Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
of ant dulosis and almost all of it has at least some empirical evidence to defend its positions. Notably, that nearly all raiding ant species have is an adaptation of their mandibles that make them big or piercing or both. It makes sense that the very thing that helps an ant be a better raider makes that same ant depend on others. The parasite loses the use of its mandibles for actual work. On this and other evidence, some propose that
predation
Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
is the precursor to slavery.
It has also been suggested that colony multiplication by adoption and budding followed by temporary parasitism can lead to dulosis. While observations indicate that colony multiplication can lead to temporary parasitism, it is apparently very questionable that temporary parasitism leads to dulosis.
In addition, about 100 years ago,
Carlo Emery
Carlo Emery (25 October 1848, Naples – 11 May 1925) was an Italian entomologist. He is remembered for Emery's rule, which states that insect social parasitism (biology), social parasites are often closely related to their hosts.
Early in hi ...
observed that, "The slave-making temporarily and permanently parasitic ants originate from closely related forms which serve them as host."
Jurgen Heinze did a series of experiments that prove this observation, known today as
Emery's Rule. Rather than just relying solely on morphological cues, he used
enzyme
An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
gel
electrophoresis
Electrophoresis is the motion of charged dispersed particles or dissolved charged molecules relative to a fluid under the influence of a spatially uniform electric field. As a rule, these are zwitterions with a positive or negative net ch ...
assays to create a detailed
phylogenetic
In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
picture of host and parasite connections.
His experimental results support a loose version of Emery’s rule. On one end of the spectrum, they saw near identical
electromorphs between species in most host and parasite pairs studied. However, it also appears that at least one example does not support Emery’s rule in that (''
Leptothorax paraxenus'') differed from its host in several electromorphs.
The selective forces involved in the evolution of social parasitism are not fully understood, and it is unlikely that one model fits all relationships.
"Any
hypothesis
A hypothesis (: hypotheses) is a proposed explanation for a phenomenon. A scientific hypothesis must be based on observations and make a testable and reproducible prediction about reality, in a process beginning with an educated guess o ...
explaining the evolution of slavery in ''Polyergus'' must account both for the origin of group raiding with brood capture, and for non-independent colony foundation by queens".
Topoff gives an adequate hypothesis to the evolution of ''Polyergus'' by integrating three processes: queen takeover, olfactory imprinting, and territorial fighting.
Here are his main points. A free-living ancestor of Polyergus, that is a
scavenger
Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
, would band together with others for intraspecific contest. Occasionally a queen invaded colonies of ''Formica''. Initially the queen drove off the ''Formica'' queen and workers and appropriated the ''Formica'' pupae (he gives as example the queens of ''
Formica wheeleri'' that do this now). The killing of the ''Formica'' queen and adoption by the workers evolved from this. His previous work (which was presented earlier) shows what would have evolved for adaptation to occur: "(1) sharp mandibles for killing the ''Formica'' queen, (2) a pheromone that reduces aggression from resident workers, (3) a tendency to hold onto the dead ''Formica'' queen long enough to absorb her odors...".
The next step was
olfactory
The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste.
In humans, it ...
imprinting between the two species, a consequence of living one’s life from start to finish in the presence of both species. ''Polyergus''’ ancestor then forages and encounters a colony of the same species. Again, they recruit nestmates to a territorial raid. In a brilliant insight, Topoff suggest this trait explains the raiding behavior. The ''Polyergus'' worker, raised by ''Formica'' as conspecifics, "identifies individual of ''Formica'' as belonging to its own species, another territorial raid is incited. Thus from the standpoint of ''Polyergus''... slave raids are equivalent to territorial raids".
The captured brood is imprinted as well and they become workers in the original nest. As time went on "Our ancestral ''Polyergus'' could easily slide in the direction of facultative parasitism".
Eventually, ''Polyergus'' ancestors lost the ability to take care of themselves and become the inquiline ''Polyergus'' we see today.
References
{{Taxonbar, from=Q1945673
Formicinae
Slave-making ants
Endemic insects of the United States
Insects described in 1893
Hymenoptera of North America
Taxonomy articles created by Polbot