Sociality is the degree to which individuals in ananimalpopulation tend to associate insocial groups (for which, the desire or inclination is known asgregariousness) and form cooperativesocieties.
This wasp behaviour demonstrates the most fundamental characteristic of animal sociality:parental investment. Parental investment is any expenditure ofresources (time, energy,social capital) to benefit one'soffspring. Parental investment detracts from a parent's capacity to invest in future reproduction and aid tokin (including other offspring). An animal that cares for its young but shows no other sociality traits is said to besubsocial.
One characteristic of social animals is the relatively high degree of cognitive ability. Social mammal predators such asspotted hyena andlion have been found to be better than non-social predators such asleopard andtiger at solving problems that require the use of innovation.[3]
Solitary animals such as thejaguar do not associate except forcourtship andmating.[4] 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 bepresocial.[5] Although presocial species are much more common than eusocial species, eusocial species have disproportionately large populations.[6]
TheentomologistCharles D. Michener published a classification system for presociality in 1969, building on the earlier work ofSuzanne Batra (who coined the wordseusocial andquasisocial in 1966).[7][8] Michener used these terms in his study of bees, but also saw a need for additional classifications:subsocial,communal, andsemisocial. In his use of these words, he did not generalize beyondinsects.E. O. Wilson later refined Batra's definition ofquasisocial.[9][10]
Subsociality is common in the animal kingdom. In subsocialtaxa, 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 besolitary but social. See Wilson (1971)[9] for definitions and further sub-classes of varieties of subsociality. Choe & Crespi (1997)[11] and Costa (2006)[12] 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 inbold italics on aphylogenetic tree of theNeoptera (note that many non-subsocial groups are omitted):[13]
Solitary-but-social animals forage separately, but some individuals sleep in the same location or share nests. Thehome 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. However, this is opposite amongcassowaries, for example. Amongprimates, this form of social organization is most common among the nocturnalstrepsirrhine species andtarsiers. Solitary-but-social species includemouse lemurs,lorises, andorangutans.[63]
Some individualcetaceans 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 includingbottlenose dolphin,common dolphin,striped dolphin,beluga,Risso's dolphin, andorca. Notable individuals includePelorus Jack (1888–1912),Tião (1994–1995), andFungie (1983–2020). At least 32 solitary-sociable dolphins were recorded between 2008 and 2019.[64]
Sociobiologists place communal, quasisocial, and semisocial animals into a meta-class: theparasocial. The two commonalities of parasocial taxa are the exhibition of parental investment, and socialization in a single,cooperative dwelling.[5]
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 ofbrood care. (This has been observed in someHymenoptera and spider taxa,[65] as well as in some otherinvertebrates.)[5] 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.
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, and wasps) andBlattodea (infraorderIsoptera, termites), but someColeoptera (such as the beetleAustroplatypus incompertus),Hemiptera (bugs such asPemphigus spyrothecae), andThysanoptera (thrips) are described as eusocial. Eusocial species that lack this criterion of morphological caste differentiation are said to beprimitively eusocial.[5]
Two potential examples of primitively eusocial mammals are thenaked mole-rat and theDamaraland mole-rat (Heterocephalus glaber andFukomys damarensis, respectively).[66] Both species arediploid and highlyinbred, 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 regularinbreeding avoidance, mole rats sometimes outbreed and establish new colonies when resources are sufficient.[67]
Eusociality has arisen among somecrustaceans that live in groups in a restricted area.Synalpheus regalis are snapping shrimp that rely on fortress defense. They live in groups of closely related individuals, amidst tropicalreefs andsponges.[68] 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.[69]
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