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''Rossomyrmex'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
slave-making ant Slave-making ants are brood parasites that capture broods of other ant species to increase the worker force of their colony. After emerging in the slave-maker nest, slave workers work as if they were in their own colony, while parasite workers onl ...
in the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Formicinae The Formicinae are a subfamily within the Formicidae containing ants of moderate evolutionary development. Formicines retain some primitive features, such as the presence of cocoons around pupae, the presence of ocelli in workers, and little ...
. The genus consists of four
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, each with a single
host A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places * Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman * Michel Host ...
from the genus ''
Proformica ''Proformica'' is a genus of ants in the subfamily Formicinae. The genus is known from the Palearctic realm, from Mongolia through Central Asia to Spain. Colonies are small, generally containing a few hundred individuals, with a single queen ( ...
'', and has a very wide range of distribution from
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 ...
to southeastern
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
, from huge extended plains to the top of high
mountains A mountain is an elevated portion of the Earth's crust, generally with steep sides that show significant exposed bedrock. Although definitions vary, a mountain may differ from a plateau in having a limited summit area, and is usually higher th ...
.


Species

* ''
Rossomyrmex anatolicus ''Rossomyrmex'' is a genus of slave-making ant in the subfamily Formicinae. The genus consists of four species, each with a single host from the genus ''Proformica'', and has a very wide range of distribution from China to southeastern Spain, f ...
'' Tinaut, 2007 –
Turkey Turkey ( tr, Türkiye ), officially the Republic of Türkiye ( tr, Türkiye Cumhuriyeti, links=no ), is a list of transcontinental countries, transcontinental country located mainly on the Anatolia, Anatolian Peninsula in Western Asia, with ...
* ''
Rossomyrmex minuchae ''Rossomyrmex minuchae'' is a species of slave-making ant in the subfamily Formicinae. It is native to Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto ...
'' Tinaut, 1981 – Spain * ''
Rossomyrmex proformicarum ''Rossomyrmex proformicarum'' is a species of slave-making ant in the subfamily Formicinae. It is native to Russia Russia (, , ), or the Russian Federation, is a transcontinental country spanning Eastern Europe and Northern Asia. It ...
'' Arnol'di, 1928 –
Caucasus The Caucasus () or Caucasia (), is a region between the Black Sea and the Caspian Sea, mainly comprising Armenia, Azerbaijan, Georgia, and parts of Southern Russia. The Caucasus Mountains, including the Greater Caucasus range, have historically ...
and
Volga The Volga (; russian: Во́лга, a=Ru-Волга.ogg, p=ˈvoɫɡə) is the List of rivers of Europe#Rivers of Europe by length, longest river in Europe. Situated in Russia, it flows through Central Russia to Southern Russia and into the Cas ...
plains,
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 ...
* ''
Rossomyrmex quandratinodum ''Rossomyrmex'' is a genus of slave-making ant in the subfamily Formicinae. The genus consists of four species, each with a single host from the genus ''Proformica'', and has a very wide range of distribution from China to southeastern Spain, fr ...
'' Xia & Zheng, 1995 –
Kazakhstan Kazakhstan, officially the Republic of Kazakhstan, is a transcontinental country located mainly in Central Asia and partly in Eastern Europe. It borders Russia to the north and west, China to the east, Kyrgyzstan to the southeast, Uzbeki ...
and China


Distribution

The Asian parasite-host pairs live mostly in extended plains whereas the Spanish pair ''R. minuchae''–''P. longiseta'' inhabits the top of three high mountains in southern Spain. Despite this apparent difference in habitat (extended plains versus high mountains), the
abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...
conditions are quite similar and are consistent with a typical arid
steppe In physical geography, a steppe () is an ecoregion characterized by grassland plains without trees apart from those near rivers and lakes. Steppe biomes may include: * the montane grasslands and shrublands biome * the temperate grasslands, ...
. However, the main difference comes from the fact that the Spanish populations are small and are geographically isolated from each other.


Slave-making

''Rossomyrmex'' is one of two
obligate {{wiktionary, obligate As an adjective, obligate means "by necessity" (antonym ''facultative'') and is used mainly in biology in phrases such as: * Obligate aerobe, an organism that cannot survive without oxygen * Obligate anaerobe, an organism that ...
slave-making genera in the subfamily Formicinae, the other being ''
Polyergus ''Polyergus'' is a small genus of ants with 14 described species. They are also referred to by the names "Slave-raiding ants" or "Amazon ants". They are characterized by their habit of raiding nests (of Formica) for workers. Reproduction ''Po ...
''. Both genera evolved slave-making behavior independently and are close
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
relatives. However, they are more closely related to the genera they parasitize than to each other, and ''Rossomyrmex'' is more closely related to ''
Cataglyphis ''Cataglyphis'' is a genus of ant, desert ants, in the subfamily Formicinae. Its most famous species is ''C. bicolor'', the Sahara Desert ant, which runs on hot sand to find insects that died of heat exhaustion, and can, like other several oth ...
'' than to ''Polyergus''.


Parasite–host pairs

Each
parasite 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 ...
species has a single host from the genus ''Proformica'', thus forming unique coevolving pairs: *''Rossomyrmex proformicarum''–''
Proformica epinotalis ''Proformica'' is a genus of ants in the subfamily Formicinae. The genus is known from the Palearctic realm, from Mongolia through Central Asia to Spain. Colonies are small, generally containing a few hundred individuals, with a single queen ( m ...
'' *''Rossomyrmex quandratinodum''–''Proformica'' sp. *''Rossomyrmex anatolicus''–''
Proformica korbi ''Proformica'' is a genus of ants in the subfamily Formicinae. The genus is known from the Palearctic realm, from Mongolia through Central Asia to Spain. Colonies are small, generally containing a few hundred individuals, with a single queen ( m ...
'' *''Rossomyrmex minuchae''–''
Proformica longiseta ''Proformica'' is a genus of ants in the subfamily Formicinae. The genus is known from the Palearctic realm, from Mongolia through Central Asia to Spain. Colonies are small, generally containing a few hundred individuals, with a single queen ( m ...
''


Raids

Parasitized nests need to replenish the host workers periodically, and this is done by raiding. The standard process is that after finding a new host nest to invade, the parasite worker marks the way to its nest with
pheromones 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 ...
, and afterward fellow slave-makers begin to attract in a few seconds. Then they go quickly to the targeted host nest, attack it, and carry as many
larvae 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 ...
and
pupae 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 ...
as possible and return to their nest following the same trail marked by the pheromone. Workers of the attacked nest can fight or flee although, in ''Proformica,'' the most common behavior is flight probably because hosts always lose fights. ''Rossomyrmex'' is the only reported slave-maker that exclusively uses adult transport and single recruitment chain instead of pheromones during raids. This behavior is probably constrained by the arid habitat: raids take place in early summer when soil surface temperature can reach up to 30 °C, a temperature for which pheromones would quickly evaporate. This condition imposes that ''Rossomyrmex'' raids appear as less efficient than those carried out with pheromones; this, together with the usually flee behavior of the ''Proformica'' hosts permits the survival of several attacked nests. Finally, another important difference in the raiding behavior of ''Rossomyrmex'' is that the return to the parasite nest with the robbed brood takes place on the following day of the assault instead of later in the same day.


Reproduction

The reproductive behavior of slave-making ants usually consists in synchronous emergence of sexuals followed by a
nuptial flight Nuptial flight is an important phase in the reproduction of most ant, termite, and some bee species. It is also observed in some fly species, such as ''Rhamphomyia longicauda''. During the flight, virgin queens mate with males and then land to ...
and the invasion of a host nest, but also in some cases females display a
mating call A mating call is the auditory signal used by animals to attract mates. It can occur in males or females, but literature is abundantly favored toward researching mating calls in females. In addition, mating calls are often the subject of mate choic ...
around the natal nest to attract males and immediately after mating search for a host nest to usurp. However, the reproductive strategy of ''Rossomyrmex'' greatly differs from the one described above. Males and females emerge from the natal nest at a different time during the day and males always fly away short after their emergence. Virgin females of ''Rossomyrmex'' show a typical mating call behavior near the natal nest but due to the scarce number of nests and that sexuals are not produced every year in all nests, some females remain virgin and cannot produce new nests despite performing sexual calling chorus for several days. When a male arrives at a female-calling nest, he will mate to as many females as possible, being one of the few cases known of polygamous males in ants, especially when mating occurs out of the nest. In contrast, females are strictly monandrous although there are some reported cases of multiply mated
queens Queens is a borough of New York City, coextensive with Queens County, in the U.S. state of New York. Located on Long Island, it is the largest New York City borough by area. It is bordered by the borough of Brooklyn at the western tip of Long ...
. Females recently mated always run to hide in their natal nest after the first copulation and do not seek for subsequent mating. This reproductive behavior seems to be constrained by the low production of sexuals, especially males (which gives advantage to female-calling behavior rather than nuptial flights and multiple mating by males). Newly mated queens search for a host nest to invade and they are unchallenged by host workers and queens thanks to the repellent effect of the
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 ...
that they have highly inflated before the usurpation. After taking over the host nest by killing the resident queens, the size of this gland decreases. This strategy to invade a host nest contrasts with other extended strategy consisting in newly mated queens embarking in a slave raid with workers, which would facilitate the penetration of the host nest immersed in chaos. In the ''Proformica''-''Rossomyrmex'' system, dispersal ability is quite different for host and parasite species. ''Proformica'' is generally
polygynous Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
(multiple queen colonies) with wingless queens that found new nests by budding; therefore they are likely to show restricted dispersal and strong population structure, while ''Rossomyrmex'' is monogynous (single queen colonies), with both sexes winged and show independent colony founding.


References

* * * * * * * * * *


External links

* {{Taxonbar, from=Q1935313 Formicinae Ant genera Hymenoptera of Asia Hymenoptera of Europe Slave-making ants Taxonomy articles created by Polbot