![Canis lupus pack surrounding Bison](https://upload.wikimedia.org/wikipedia/commons/4/45/Canis_lupus_pack_surrounding_Bison.jpg)
Sociality is the degree to which individuals in an
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
population
Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...
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 varieties ...
s (gregariousness) and form cooperative
societies.
Sociality is a survival response to
evolutionary pressures.
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 suborder. Th ...
stays near her
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 in the nest,
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 ...
s 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
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Vespidae.
This wasp behaviour evidences the most fundamental characteristic of animal sociality:
parental investment
Parental investment, in evolutionary biology and evolutionary psychology, is any parental expenditure (e.g. time, energy, resources) that benefits offspring.Clutton-Brock, T.H. 1991. ''The Evolution of Parental Care''. Princeton, NJ: Princeton ...
. Parental investment is any expenditure of
resource
Resource refers to all the materials available in our environment which are technologically accessible, economically feasible and culturally sustainable and help us to satisfy our needs and wants. Resources can broadly be classified upon their ...
s (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 (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 als ...
division of labor
The division of labour is the separation of the tasks in any economic system or organisation so that participants may specialise (specialisation). Individuals, organizations, and nations are endowed with, or acquire specialised capabilities, and ...
, 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 cultura ...
.
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 th ...
do not associate except for
courtship
Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...
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 arach ...
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
insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s.
E. O. Wilson 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, 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
A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
of the
Neoptera (note that many non-subsocial groups are omitted):
Solitary but social
![Microcebus murinus -Artis Zoo, Amsterdam, Netherlands-8c](https://upload.wikimedia.org/wikipedia/commons/e/ed/Microcebus_murinus_-Artis_Zoo%2C_Amsterdam%2C_Netherlands-8c.jpg)
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
primate
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s, this form of social organization is most common among the nocturnal
strepsirrhine 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 Southeast ...
s. Solitary-but-social species include
mouse lemurs,
lorises, and
orangutan
Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s.
Some individual
cetacean
Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
s 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
Bottlenose dolphins are aquatic mammals in the genus ''Tursiops.'' They are common, cosmopolitan members of the family Delphinidae, the family of oceanic dolphins. Molecular studies show the genus definitively contains two species: the common ...
,
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 (biology), family, Delphinidae.
Taxonomy
The stripe ...
,
beluga,
Risso's dolphin, and
orca
The orca or killer whale (''Orcinus orca'') is a toothed whale belonging to the oceanic dolphin family, of which it is the largest member. It is the only Extant taxon, extant species in the genus ''Orcinus'' and is recognizable by its black ...
. Notable individuals include
Pelorus Jack (1888–1912),
Tião (1994–1995), and
Fungie (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
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 br ...
. (This has been observed in some
Hymenoptera
Hymenoptera is a large order (biology), order of insects, comprising the sawfly, 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 Par ...
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
![Natural Beehive and Honeycombs](https://upload.wikimedia.org/wikipedia/commons/9/92/Natural_Beehive_and_Honeycombs.jpg)
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,
bee
Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyly, monophyletic lineage within the ...
s,
sawflies, and wasps) and
Blattodea
Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence suggests they evolved from within the cockroach lineage, cladistically ...
(infraorder
Isoptera
Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
, termites), but some
Coleoptera
Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
(such as the beetle ''
Austroplatypus incompertus''),
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. ...
(bugs such as ''
Pemphigus spyrothecae
''Pemphigus spyrothecae'', or the poplar spiral gall aphid, is a social insect which exhibits apparent altruistic behaviors. The aphids form galls and act as colony defenders, at times sacrificing their own lives to do so. It has been shown that ...
''), and
Thysanoptera (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 (''Heterocephalus glaber'' and ''Fukomys damarensis'', respectively). Both species are
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
and highly
inbred, 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
Inbreeding avoidance, or the inbreeding avoidance hypothesis, is a concept in evolutionary biology that refers to the prevention of the deleterious effects of inbreeding. Animals only rarely exhibit inbreeding avoidance. The 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 ...
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
reef
A reef is a ridge or shoal of rock, coral or similar relatively stable material, lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic processes— deposition of sand, wave erosion planing down rock out ...
s 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 t ...
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 and
Bert Hölldobler controversially claimed in 2005 that
human
Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
s 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