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Sociality is the degree to which individuals in an animal population tend to associate in
social group In the social sciences, a social group can be defined as two or more people who interact with one another, share similar characteristics, and collectively have a sense of unity. Regardless, social groups come in a myriad of sizes and varietie ...
s (gregariousness) and form cooperative
societies A society is a group of individuals involved in persistent social interaction, or a large social group sharing the same spatial or social territory, typically subject to the same political authority and dominant cultural expectations. Societ ...
. Sociality is a survival response to
evolutionary pressure Any cause that reduces or increases reproductive success in a portion of a population potentially exerts evolutionary pressure, selective pressure or selection pressure, driving natural selection. It is a quantitative description of the amount of ...
s. For example, when a mother
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborde ...
stays near her larvae in the nest, parasites are less likely to eat the larvae. Biologists suspect that pressures from parasites and other
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s selected this behavior in wasps of the family
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as '' Polistes fuscatus'', '' Vespa orientalis'', and '' Vespula germanica'') and many solitary wasps. ...
. This wasp behaviour evidences the most fundamental characteristic of animal sociality: parental investment. Parental investment is any expenditure of resources (time, energy,
social capital Social capital is "the networks of relationships among people who live and work in a particular society, enabling that society to function effectively". It involves the effective functioning of social groups through interpersonal relationships ...
) to benefit one's
offspring In biology, offspring are the young creation of living organisms, produced either by a single organism or, in the case of sexual reproduction, two organisms. Collective offspring may be known as a brood or progeny in a more general way. This ca ...
. Parental investment detracts from a parent's capacity to invest in future reproduction and aid to
kin __NOTOC__ Kin usually refers to kinship and family. Kin or KIN may also refer to: Culture and religion * Otherkin, people who identify as not entirely human * Kinism, a white supremacist religious movement * Kinh, the majority ethnic group o ...
(including other offspring). An animal that cares for its young but shows no other sociality traits is said to be ''subsocial''. An animal that exhibits a high degree of sociality is called a ''social animal''. The highest degree of sociality recognized by sociobiologists is '' eusociality''. A eusocial
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
is one that exhibits overlapping adult generations,
reproductive The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are a ...
division of labor, cooperative care of young, and—in the most refined cases—a biological
caste system Caste is a form of social stratification characterised by endogamy, hereditary transmission of a style of life which often includes an occupation, ritual status in a hierarchy, and customary social interaction and exclusion based on cultural ...
.


Presociality

Solitary animals such as the
jaguar The jaguar (''Panthera onca'') is a large cat species and the only living member of the genus ''Panthera'' native to the Americas. With a body length of up to and a weight of up to , it is the largest cat species in the Americas and the thi ...
do not associate except for courtship and mating. If an animal taxon shows a degree of sociality beyond courtship and mating, but lacks any of the characteristics of eusociality, it is said to be ''presocial''. Although presocial species are much more common than eusocial species, eusocial species have disproportionately large populations. The
entomologist Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as ar ...
Charles D. Michener published a classification system for presociality in 1969, building on the earlier work of
Suzanne Batra Suzanne Wellington Tubby Batra (born December 15, 1937) is an American entomologist best known for her work on the classification of insect societies and for coining the term eusociality. Batra was born in New York City where her father Roger W ...
(who coined the words ''eusocial'' and ''quasisocial'' in 1966). Michener used these terms in his study of bees, but also saw a need for additional classifications: ''subsocial'', ''communal'', and ''semisocial''. In his use of these words, he did not generalize beyond insects.
E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, entomologist and writer. According to David Attenborough, Wilson was the world's leading expert in his specialty of myrmecology, the study of an ...
later refined Batra's definition of ''quasisocial''. Entry is linked to entries on each of the other terms, as Subsocial, Quasisocial, etc.


Subsociality

Subsociality is common in the animal kingdom. In subsocial
taxa In biology, a taxon (back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular na ...
, parents care for their young for some length of time. Even if the period of care is very short, the animal is still described as subsocial. If adult animals associate with other adults, they are not called subsocial, but are ranked in some other classification according to their social behaviours. If occasionally associating or nesting with other adults is a taxon's most social behaviour, then members of those populations are said to be ''solitary but social''. See Wilson (1971) for definitions and further sub-classes of varieties of subsociality. Choe & Crespi (1997) and Costa (2006) give readable overviews. Subsociality is widely distributed among the winged insects, and has evolved independently many times. Insect groups that contain at least some subsocial species are shown in boldface on a phylogenetic tree of the
Neoptera Neoptera (Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world Ancient history is a time period from the beginning of writing and recorded human history to as fa ...
(note that many non-subsocial groups are omitted):


Solitary but social

Solitary-but-social animals forage separately, but some individuals sleep in the same location or share nests. The home ranges of females usually overlap, whereas those of males do not. Males usually do not associate with other males, and male offspring are usually evicted upon maturity. Among primates, this form of social organization is most common among the nocturnal
strepsirrhine Strepsirrhini or Strepsirhini (; ) is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. ...
species and
tarsier Tarsiers ( ) are haplorhine primates of the family Tarsiidae, which is itself the lone extant family within the infraorder Tarsiiformes. Although the group was once more widespread, all of its species living today are found in Maritime Southea ...
s. Solitary-but-social species include
mouse lemur The mouse lemurs are nocturnal lemurs of the genus ''Microcebus''. Like all lemurs, mouse lemurs are native to Madagascar. Mouse lemurs have a combined head, body and tail length of less than , making them the smallest primates (the smallest spe ...
s,
loris Loris is the common name for the strepsirrhine mammals of the subfamily Lorinae (sometimes spelled Lorisinae) in the family Lorisidae. ''Loris'' is one genus in this subfamily and includes the slender lorises, ''Nycticebus'' is the genus cont ...
es, and orangutans. Some individual cetaceans adopt a solitary but social behavior, that is, they live apart from their own species but interact with humans. This behavior has been observed in species including bottlenose dolphin,
common dolphin The common dolphin (''Delphinus delphis'') is the most abundant cetacean in the world, with a global population of about six million. Despite this fact and its vernacular name, the common dolphin is not thought of as the archetypal dolphin, with ...
,
striped dolphin The striped dolphin (''Stenella coeruleoalba'') is an extensively studied dolphin found in temperate and tropical waters of all the world's oceans. It is a member of the oceanic dolphin family, Delphinidae. Taxonomy The striped dolphin is on ...
, beluga,
Risso's dolphin Risso's dolphin (''Grampus griseus'') is a dolphin, the only species of the genus ''Grampus''. Some of the closest related species to these dolphins include: pilot whales (''Globicephala'' spp.), pygmy killer whales (''Feresa attenuata''), me ...
, and orca. Notable individuals include
Pelorus Jack Pelorus Jack ( fl. 1888 – April 1912) was a Risso's dolphin (''Grampus griseus'') that was famous for meeting and escorting ships through a stretch of water in Cook Strait, New Zealand. The animal was reported over a 24 year period, from ...
(1888–1912), Tião (1994–1995), and
Fungie Fungie (), also known as the Dingle Dolphin, was a male common bottlenose dolphin. He became separated from other wild dolphins and lived in very close contact with the people of Dingle on the southwest coast of Ireland. Wild bottlenose dol ...
(1983–2020). At least 32 solitary-sociable dolphins were recorded between 2008 and 2019.


Parasociality

Sociobiologists place communal, quasisocial, and semisocial animals into a meta-class: the ''parasocial''. The two commonalities of parasocial taxa are the exhibition of parental investment, and socialization in a single, cooperative dwelling. Communal, quasisocial, and semisocial groups differ in a few ways. In a communal group, adults cohabit in a single nest site, but they each care for their own young. Quasisocial animals cohabit, but they also share the responsibilities of brood care. (This has been observed in some
Hymenoptera Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typic ...
and spider taxa, as well as in some other
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s.) A semisocial population has the features of communal and quasisocial populations, but they also have a biological caste system that delegates labor according to whether or not an individual is able to reproduce. Beyond parasociality is eusociality. Eusocial insect societies have all the characteristics of a semisocial one, except overlapping generations of adults cohabit and share in the care of young. This means that more than one adult generation is alive at the same time, and that the older generations also care for the newest offspring.


Eusociality

Eusocial societies have overlapping adult generations, cooperative care of young, and division of reproductive labor. When organisms in a species are born with physical characteristics specific to a caste which never changes throughout their lives, this exemplifies the highest acknowledged degree of sociality. Eusociality has evolved in several orders of insects. Common examples of eusociality are from Hymenoptera ( ants, bees,
sawflies Sawflies are the insects of the suborder Symphyta within the order Hymenoptera, alongside ants, bees, and wasps. The common name comes from the saw-like appearance of the ovipositor, which the females use to cut into the plants where they lay ...
, and wasps) and Blattodea (infraorder Isoptera, termites), but some Coleoptera (such as the beetle ''
Austroplatypus incompertus ''Austroplatypus incompertus'' is a species of ambrosia beetle belonging to the true weevil family, native to Australia, with a verified distribution in New South Wales and Victoria. It forms colonies in the heartwood of ''Eucalyptus'' trees ...
''), Hemiptera (bugs such as '' Pemphigus spyrothecae''), and
Thysanoptera Thrips ( order Thysanoptera) are minute (mostly long or less), slender insects with fringed wings and unique asymmetrical mouthparts. Different thrips species feed mostly on plants by puncturing and sucking up the contents, although a few are ...
(thrips) are described as eusocial. Eusocial species that lack this criterion of morphological caste differentiation are said to be ''primitively eusocial''. Two potential examples of primitively eusocial mammals are the
naked mole-rat The naked mole-rat (''Heterocephalus glaber''), also known as the sand puppy, is a burrowing rodent native to the Horn of Africa and parts of Kenya, notably in Somali regions. It is closely related to the blesmols and is the only species in th ...
and the
Damaraland mole-rat The Damaraland mole-rat (''Fukomys damarensis''), Damara mole rat or Damaraland blesmol, is a burrowing rodent found in southern Africa. Along with the smaller, less hairy, naked mole rat, it is a species of eusocial mammal. Description Like ot ...
(''Heterocephalus glaber'' and ''Fukomys damarensis'', respectively). Both species are diploid and highly
inbred Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders and o ...
, and they aid in raising their siblings and relatives, all of whom are born from a single reproductive queen; they usually live in harsh or limiting environments. A study conducted by O'Riain and Faulkes in 2008 suggests that, due to regular inbreeding avoidance, mole rats sometimes outbreed and establish new colonies when resources are sufficient. Eusociality has arisen among some
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group c ...
s that live in groups in a restricted area. ''
Synalpheus regalis ''Synalpheus regalis'' is a species of snapping shrimp that commonly live in sponges in the coral reefs along the tropical West Atlantic. They form a prominent component of the diverse marine cryptofauna of the region. For the span of their en ...
'' are snapping shrimp that rely on fortress defense. They live in groups of closely related individuals, amidst tropical reefs and
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through ...
s. Each group has one breeding female; she is protected by a large number of male defenders who are armed with enlarged snapping claws. As with other eusocial societies, there is a single shared living space for the colony members, and the non-breeding members act to defend it.


Human eusociality

E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, entomologist and writer. According to David Attenborough, Wilson was the world's leading expert in his specialty of myrmecology, the study of an ...
and Bert Hölldobler controversially claimed in 2005 that humans exhibit sufficient sociality to be counted as a eusocial species, and that this enabled them to enjoy spectacular ecological success and dominance over ecological competitors.


See also


References

{{Sociobiology Behavioral ecology Ethology Sociobiology