''Tetraponera'' 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 ...

ants 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 Creta ...

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 ...

Pseudomyrmecinae Pseudomyrmecinae is a small subfamily of ants containing only three genera of slender, large-eyed arboreal ants, predominantly tropical or subtropical in distribution. In the course of adapting to arboreal conditions (unlike the predominantly ...

that are commonly known as slender ants and are characterized by their

arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose nu ...

nature and slender bodies. The 96 described

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 ...

of ''Tetraponera'' all of which live in hollow structures of plants and trees, such as thorns or branches; these hosts are known as

myrmecophyte Myrmecophytes (; literally "ant-plant") are plants that live in a mutualistic association with a colony of ants. There are over 100 different genera of myrmecophytes. These plants possess structural adaptations that provide ants with food and ...

s. ''Tetraponera'' species are closely related to the New World genus of ants ''

Pseudomyrmex ''Pseudomyrmex'' is a genus of stinging, wasp-like ants in the subfamily Pseudomyrmecinae. They are large-eyed, slender ants, found mainly in tropical and subtropical regions of the New World. Distribution and habitat ''Pseudomyrmex'' is predom ...

'', but differ in their relationships with host plants.

Mutualisms and behaviour

''Tetraponera'' species are generally defined by the myrmecophytes they inhabit and the mutualistic relationship they share. These host plants always have hollow thorns or branches in which the

ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...

s can live and form a

colony In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the ''metropole, metropolit ...

. Also, the myrmecophytes provide energy rich food sources such as extrafloral nectar and/or food bodies. All ''Tetraponera'' species have gut

symbiont Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...

s that allow them to digest

amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...

-deficient food provided by their host plants; these

gut bacteria Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut ...

are especially important for the species that only survive on the myrmecophyte-provided foods. All ''Tetraponera'' species provide protection for their host plants through aggressive nature towards other insects and trimming leaves/branches of neighbouring plants. Living in hollow structures of the plants allows the ants to detect vibrations when larger insects land on the plant, or workers on patrol visually detect smaller intruders. Once detected, sophisticated

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 ...

systems allow the ants to quickly outnumber and overpower any invaders. Most insect invaders are killed and discarded by ''Tetraponera'' workers such as

caterpillars Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Symp ...

and

aphids Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...

, but some are killed and consumed. The insects that take the most time and effort to kill are generally consumed; these are mostly

katydids Insects in the family Tettigoniidae are commonly called katydids (especially in North America), or bush crickets. They have previously been known as "long-horned grasshoppers". More than 8,000 species are known. Part of the suborder Ensifera, t ...

or leaf beetles. As well as assaulting and killing insects that attack their host plant, ''Tetraponera'' ants will attack any

mammals Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...

that present a threat. Inflicting the mammals with painful stings will usually deter them from attacking the plant. In some arboreal ant species, not just ''Tetraponera'', a third partner in ant-myrmecophyte mutualisms is

hemiptera Hemiptera (; ) is an order (biology), order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, Reduviidae, assassin bugs, Cimex, bed bugs, and shield bugs. ...

n trophobionts. These insects provide a possible third source of food for the ants, in return the ants feed and “nurture” the Hemiptera inside their

colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...

Distribution

''Tetraponera'' species are found commonly in the warmer regions of

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 ...

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...

, and

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

. Different species are associated with different plant species. The most common myrmecophytes for ''Tetraponera'' are

acacias ''Acacia'', commonly known as the wattles or acacias, is a large genus of shrubs and trees in the subfamily Mimosoideae of the pea family Fabaceae. Initially, it comprised a group of plant species native to Africa and Australasia. The genus nam ...

, but the wide variety of hosts for this genus include

bamboo Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, bu ...

s and

lianas A liana is a long- stemmed, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the canopy in search of direct sunlight. The word ''liana'' does not refer to a t ...

. ''Tetraponera'' ants trim neighbouring plants to prevent any intrusion of other ants or caterpillars from those plants and to reduce resource competition for their

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 ...

. ''Tetraponera'' species only leave their myrmecophyte to start a new colony on a different host; a

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 ...

and a number of

workers The workforce or labour force is a concept referring to the pool of human beings either in employment or in unemployment. It is generally used to describe those working for a single company or industry, but can also apply to a geographic reg ...

leave the old host plant to allow the colony to continue to expand.

Lifecycle and castes

Tetraponera natalensis, werkerkaste, Pretoria

''Tetraponera'', like most ants, has one or a few queens that are the only females to

reproduce Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. Reproduction is a fundamental feature of all known life; each individual org ...

in a colony. The sterile workers are all females that forage for food and defend the colony. Males are produced only during certain times of the year and disperse to mate with virgins queens from other colonies. Since ants are

haplodiploid Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the se ...

, they can control what sex their offspring will be; an unfertilised egg will become a male, while a fertilised egg will be female. This reliably restricts the production of male alates to the species' mating season, when the winged virgin queens and males fly from their home colonies to mate and start new colonies. The eggs produced by the queen hatch into larvae which are cared for inside the colony, protected from any predators by the workers. The amount of care each female larva receives determines its fate as a worker or a new queen; all males are drones. When a new colony is formed, eggs are initially produced at a low rate, but this quickly increases in the second to fourth years, to ensure enough workers are produced to protect and provide for the growing colony.

Species

*'' T. aethiops'' Smith, 1877 *'' T. aitkenii'' (Forel, 1902) *'' T. allaborans'' (Walker, 1859) *'' T. amargina'' Xu & Chai, 2004 *'' T. ambigua'' (Emery, 1895) *'' T. andrei'' (Mayr, 1895) *'' T. anthracina'' (Santschi, 1910) *'' T. apiculata'' Ward, 2001 *'' T. atra'' Donisthorpe, 1949 *'' T. attenuata'' Smith, 1877 *'' T. avia'' Ward, 2001 *'' T. bifoveolata'' (Mayr, 1895) *'' T. binghami'' (Forel, 1902) *'' T. bita'' Ward, 2001 *'' T. brevis'' Ward, 2001 *'' T. buops'' Ward, 2001 *'' T. caffra'' (Santschi, 1914) *'' T. clypeata'' (Emery, 1886) *'' T. concava'' Xu & Chai, 2004 *'' T. conica'' Ward, 2001 *'' T. connectens'' Ward, 2001 *'' T. continua'' (Forel, 1907) *'' T. convexa'' Xu & Chai, 2004 *'' T. cortina'' Ward, 2022 *'' T. crassiuscula'' (Emery, 1900) *'' T. diana'' (Santschi, 1911) *'' T. difficilis'' (Emery, 1900) *'' T. dispar'' Ward, 2022 *'' T. elegans'' Ward, 2022 *'' T. emeryi'' (Forel, 1911) *'' T. erythraea'' (Emery, 1895) *†'' T. europaea'' Dlussky, 2009 *'' T. exactor'' Ward, 2022 *'' T. exasciata'' (Forel, 1892) *'' T. extenuata'' Ward, 2001 *'' T. fictrix'' (Forel, 1897) *'' T. furcata'' Xu & Chai, 2004 *'' T. furtiva'' Ward, 2022 *'' T. gerdae'' (Stitz, 1911) *'' T. grandidieri'' (Forel, 1891) *†'' T. groehni'' Dlussky, 2009 *'' T. hespera'' Ward, 2009 *'' T. hirsuta'' Ward, 2009 *'' T. hysterica'' (Forel, 1892) *'' T. inermis'' Ward, 2009 *'' T. insularis'' Ward, 2022 *'' T. inversinodis'' Ward, 2001 *†'' T. klebsi'' (Wheeler, 1915) *'' T. kosi'' Ward, 2022 *†'' T. lacrimarum'' (Wheeler, 1915) *'' T. laeviceps'' (Smith, 1859) *'' T. latifrons'' (Emery, 1912) *'' T. liengmei'' (Forel, 1894) *'' T. longula'' (Emery, 1895) *'' T. manangotra'' Ward, 2009 *'' T. mandibularis'' (Emery, 1895) *'' T. mayri'' (Forel, 1901) *'' T. merita'' Ward, 2009 *'' T. microcarpa'' Wu & Wang, 1990 *'' T. mimula'' Ward, 2001 *'' T. mocquerysi'' (André, 1890) *'' T. modesta'' (Smith, 1860) *'' T. morondaviensis'' (Forel, 1891) *'' T. natalensis'' (Smith, 1858) *'' T. nigra'' (Jerdon, 1851) *'' T. nitida'' (Smith, 1860) *'' T. nixa'' Ward, 2001 *'' T. nodosa'' Ward, 2001 *'' T. notabilis'' Ward, 2001 *†'' T. ocellata'' (Mayr, 1868) *†'' T. oligocenica'' (Théobald, 1937) *'' T. ophtalmica'' (Emery, 1912) *'' T. parops'' Ward, 2006 *'' T. pedana'' Ward, 2022 *'' T. penzigi'' (Mayr, 1907) *'' T. periyarensis'' Bharti & Akbar, 2014 *'' T. perlonga'' Santschi, 1928 *'' T. phragmatica'' Ward, 2006 *'' T. pilosa'' (Smith, 1858) *'' T. polita'' Ward, 2001 *'' T. protensa'' Xu & Chai, 2004 *'' T. pumila'' Ward, 2022 *'' T. punctulata'' Smith, 1877 *'' T. rakotonis'' (Forel, 1891) *'' T. redacta'' Ward, 2022 *'' T. rotula'' Ward, 2001 *'' T. rufonigra'' (Jerdon, 1851) *'' T. sahlbergii'' (Forel, 1887) *'' T. schulthessi'' (Santschi, 1915) *'' T. setosa'' Ward, 2022 *†'' T. simplex'' (Mayr, 1868) *'' T. tessmanni'' (Stitz, 1910) *'' T. tucurua'' Ward, 2001 *'' T. variegata'' (Forel, 1895) *'' T. vivax'' Ward, 2001 *'' T. volucris'' Ward, 2001

References

* Borm, S.V., A. Buschinger, J. J. Boomsma and J. Billen. 2002. Tetraponera ants have gut symbionts related to nitrogen-fixing root-nodule bacteria. Biological Sciences. 269:2023-2027. * Ward, P.S. 2001. Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera:Formicidae) in the Oriental and Australian regions Invertebrate Taxonomy. 15:589:665. * Dejean, A., J. Orivel and C. Djieto-Lordon. 2008. The plant ant Tetraponera aethiops (Pseudomyrmecinae) protects its host myrmecophyte Barteria fistulosa (Passifloraceae) through aggressiveness and predation. 93:63-69. * Australian Biological Resources and Study: Australian Faunal Directory. http://www.environment.gov.au/biodiversity/abrs/online-resources/fauna/afd/taxa/Tetraponera

External links

* * {{Taxonbar, from=Q771402 Pseudomyrmecinae Ant genera