''Formica sanguinea'', or blood-red ant, is a species of facultative slave-maker ant in the genus ''

Formica ''Formica'' is a genus of ants of the family Formicidae, 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 species of genus ''For ...

'' characterized by the ability to secrete formic acid. It ranges from

Central Central is an adjective usually referring to being in the center of some place or (mathematical) object. Central may also refer to: Directions and generalised locations * Central Africa, a region in the centre of Africa continent, also known as ...

and

Northern Europe The northern region of Europe has several definitions. A restrictive definition may describe Northern Europe as being roughly north of the southern coast of the Baltic Sea, which is about 54th parallel north, 54°N, or may be based on other g ...

through

Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...

Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...

China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...

, the

Korean Peninsula Korea ( ko, 한국, or , ) is a peninsular region in East Asia. Since 1945, it has been divided at or near the 38th parallel, with North Korea (Democratic People's Republic of Korea) comprising its northern half and South Korea (Republic o ...

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

and also the

United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...

. This species is coloured red and black with workers up to 7 mm long. A colony of ''F. sanguinea'' can live either as a free colony or as a social parasite of ''Formica'' species, most commonly ''

Formica fusca ''Formica fusca'' is a black-colored ant commonly found throughout Europe as well as parts of Southern Asia and Africa. It has the common names silky ant or dusky ant. The range within the palaearctic region extends from Portugal in the west to J ...

'', ''

Formica japonica ''Formica japonica'' is a species of ant in the genus ''Formica''. It is found in Russian (Far East The ''Far East'' was a European term to refer to the geographical regions that includes East and Southeast Asia as well as the Russian Far Ea ...

'', '' Formica hayashi'' and ''

Formica rufibarbis ''Formica rufibarbis'' is a European formicine ant of the ''Formica fusca'' group. In the classification by Auguste Forel, it is treated in the subgenus '' Serviformica''. ''F. rufibarbis'' is subject to a Species Action Plan (SAP) in Englan ...

''.

Raiding

Blood-red ants, ''F. sanguinea'', are facultative slave-makers, meaning colonies can live either alone or be

parasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...

. This allows them to be a good

model organism A model organism (often shortened to model) is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workin ...

to study the origins of brood stealing. A fertilized ''F. sanguinea''

queen Queen or QUEEN may refer to: Monarchy * Queen regnant, a female monarch of a Kingdom ** List of queens regnant * Queen consort, the wife of a reigning king * Queen dowager, the widow of a king * Queen mother, a queen dowager who is the mother ...

will enter the

nest A nest is a structure built for certain animals to hold 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 organic materia ...

of the host ant species and kill their queen. She then takes advantage of the workers who tend to her and her

brood Brood may refer to: Nature * Brood, a collective term for offspring * Brooding, the incubation of bird eggs by their parents * Bee brood, the young of a beehive * Individual broods of North American Periodical Cicadas: ** Brood X, the largest bro ...

. ''F. sanguinea'' workers will also raid nearby nests, stealing

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

e and

pupa A pupa ( la, pupa, "doll"; plural: ''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 ...

e to become future workers for ''F. sanguinea''. The raids are also not exclusively for acquiring new workers but are sometimes predation events. ''Formica sanguinea'' has not been observed to have division of labor in which certain individuals raid or forage. However, some individuals possess more

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 at a ...

compounds than others, so presumably would be more successful in during raiding. ''Formica sanguinea'' uses scouting individuals to locate the nest that will be parasitized. Once a nest has been scouted, the raid will happen. The activities that go on during a raid are composed of digging and fighting at the target nest. Both captive workers and blood-red ant raiders are observed to be carrying the brood back to the ''Formica sanguinea'' nest. If a member of the colony is killed in the raid, individuals will take that carcass back to the nest to be eaten later. Raids were also never observed on rainy or overcast days. This is thought to be because of the detriment rain might have on terrestrial conditions and the effectiveness of alarm pheromones.

Chemical defenses

''F. sanguinea'' uses

formic acid Formic acid (), systematically named methanoic acid, is the simplest carboxylic acid, and has the chemical formula HCOOH and structure . It is an important intermediate in chemical synthesis and occurs naturally, most notably in some ants. Es ...

as well as substances in the

as chemical defence. The substances in the Dufour's gland contain

hydrocarbons In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and hydrophobic, and their odors are usually weak or ex ...

such as hendecane, as well as

acetate An acetate is a salt (chemistry), salt formed by the combination of acetic acid with a base (e.g. Alkali metal, alkaline, Alkaline earth metal, earthy, Transition metal, metallic, nonmetallic or radical Radical (chemistry), base). "Acetate" als ...

s, decylacetate, and dodecylacetate. ''Formica sanguinea'' uses the formic acid and Dufour's gland substances in conjunction with each other. The hydrocarbons in Dufour’s substance serve as a wetting before the formic acid is released. Without these wetting agents, formic acid is relatively harmless to other ants. These substances dissolve the fat compounds of the

epicuticle The cuticle forms the major part of the integument of the Arthropoda. It includes most of the material of the exoskeleton of the insects, Crustacea, Arachnida, and Myriapoda. Morphology In arthropods, the integument, the external "skin", or ...

as well as enter the tracheal system to kill the opponent. Interactions between Formica sanguinea and Lomechusoides strumosus

Reproduction

Most colonies are

monogynous This is a glossary of terms used in the descriptions of ants. A B D E F G H M N O P Q R S T U W See also ...

, meaning they have only one queen. ''F. sanguinea'' are commonly

polyandrous Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wives" ...

, meaning the singular queen mates with more than one male. In studies observing this, 70% of queens mated with multiple males. ''F. sanguinea'' is observed to have higher levels of polyandry than any other ''Formica'' species. However, when colonies engage in polygyny, having more than one queen, the rates of polyandry drop. It is assumed that colonies utilize either polyandry or polygyny, to increase the

genetic diversity Genetic diversity is the total number of genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. It is dis ...

of the colony. Therefore, two different strategies can be employed. One strategy is that a female mates with many males and then disperses to start a new colony. The second strategy is for a female to mate with a fewer number of males, and stay close to the natal colony with other females who have mated with a fewer number of males. In the case of polygyny, it has been observed that there is one dominating queen who is more sexual reproductive than the others. Formica also is observed to have a paternity skew, meaning that the mated males offspring are not represented equally in the population. Whether this is because of internal selection or

sperm competition Sperm competition is the competitive process between spermatozoa of two or more different males to fertilize the same egg during sexual reproduction. Competition can occur when females have multiple potential mating partners. Greater choice and ...

is unclear.

Recognition of individuals

''F. sanguinea'' analyzes hydrocarbons on the cuticle of another individual to determine if it is an intruder. Low

intraspecific competition Intraspecific competition is an interaction in population ecology, whereby members of the same species compete for limited resources. This leads to a reduction in fitness for both individuals, but the more fit individual survives and is able to r ...

and aggressiveness is observed in this species. When members of the same species of a mixed colony were met with individuals from a pure ''F. sanguinea'' colony, they were met with aggressiveness. This is most likely because the identifying hydrocarbons change when individuals are in a mixed colony.

References

External links

* {{Taxonbar, from=Q1350298 sanguinea Slave-making ants Hymenoptera of Europe Hymenoptera of Asia Insects described in 1798