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Sociality
Sociality is the degree to which individuals in an animal
population tend to associate in social groups
(gregariousness) and form cooperative societies.

Sociality is a survival response to evolutionary pressures.[1]

For example, when a mother wasp stays near her larvae in the
nest, parasites are less likely to eat the larvae.[2] Biologists
suspect that pressures from parasites and other predators
selected this behavior in wasps of the family Vespidae.

This wasp behaviour evidences the most fundamental

characteristic of animal sociality: parental investment.
Gray wolves hunting in a presocial pack
Parental investment is any expenditure of resources (time,
encircle an American bison.
energy, social capital) to benefit one's offspring. 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 is one that exhibits overlapping
adult generations, reproductive division of labor, cooperative care of young, and—in the most refined
cases—a biological caste system.

Contents
Presociality
Subsociality
Solitary but social
Parasociality
Eusociality
Human eusociality
See also
References

Presociality
Solitary animals such as the jaguar do not associate except for courtship and mating.[3] 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.[4] Although presocial species are much more common than
eusocial species, eusocial species have disproportionately large populations.[5]

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The entomologist Charles D. Michener published a classification system for presociality in 1969,
building on the earlier work of Suzanne Batra (who coined the words eusocial and quasisocial in
1966).[6][7] 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 later refined Batra's definition of quasisocial.[8][9]

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)[8] for definitions and further sub-classes of varieties of
subsociality. Choe & Crespi (1997)[10] and Costa (2006)[11] 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 (note that many non-subsocial groups are omitted):[12]
Neoptera  
    Idioprothoraca
  Embioptera (webspinners)[13][14]

  [15]
  Blattodea (cockroaches, inc. eusocial termites)
Dictyoptera
 
 
  Mantodea (mantises)
 
 
Rhipineoptera
 
  [16]
  Orthoptera (grasshoppers, crickets)
 
 
  [17][18][19]
  Dermaptera (earwigs)
Eumetabola Parametabola
     
  Zoraptera (angel insects)[20]
Paraneoptera Condylognatha
     
  Thysanoptera (thrips)[21]
Hemiptera (bugs)
many families[30][31] Membracidae
  (treehoppers, thorn
 
bugs)[22][23]
 
 
  Pentatomidae (shield
  bugs)[24]

  
   

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Reduviidae
(predatory
bugs)[25][26][27]

  Tingidae (lace
  bugs)[28][29]

 
  Psocoptera (bark lice)[32]

  Staphylinidae (rove
  beetles)[33][34][35][36][37][38][39]
 
 
  [40]
  Silphidae (carrion beetles)
 
 
  [41][42][43][44]
  Passalidae (bessbugs)
 
 
  [45][46]
  Scarabaeidae (scarabs)
Coleoptera[59]
    [47][48]
    Tenebrionidae (leaf/flower beetles)
 

  [49]
    Erotylidae (pleasing fungus beetles)
 
 
 
  Chrysomelidae (leaf
Endopterygota   beetles)[50][51][52][53][54][55][56][57][58]
 

 
  Raphidioptera (snakeflies)
Neuropteroidea
 
 
  Neuroptera (lacewings, alderflies, and allies)

 
  Antliophora (true flies, scorpionflies, fleas)

   
    Trichoptera (caddisflies)
 
   
    [60]
  Lepidoptera (butterflies and moths)

[61] (apart from

  Hymenoptera (sawflies, wasps, ants, bees)
  eusocial species)

Solitary but social

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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 species and tarsiers. Solitary-but-social species
include mouse lemurs, lorises, and orangutans.[62]
The mouse lemur is a nocturnal,
Some individual cetaceans adopt a solitary but social behavior,
solitary-but-social lemur native to
that is, they live apart from their own species but interact with
Madagascar.
humans. This behavior has been observed in species including
bottlenose dolphin, common dolphin, striped dolphin, beluga,
Risso's dolphin, and orca. 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.[63]

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.[4]

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 and
spider taxa,[64] as well as in some other invertebrates.)[4] 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, and wasps) and Blattodea (infraorder Isoptera,
termites), but some Coleoptera (such as the beetle Austroplatypus Giant honey bees cover the
incompertus), Hemiptera (bugs such as Pemphigus spyrothecae), honeycomb of their nest.
and Thysanoptera (thrips) are described as eusocial. Eusocial
species that lack this criterion of morphological caste
differentiation are said to be primitively eusocial.[4]
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Two potential examples of primitively eusocial mammals are the naked mole-rat and the Damaraland
mole-rat (Heterocephalus glaber and Fukomys damarensis, respectively).[65] Both species are diploid
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. However, 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.[66]

Eusociality has arisen among some crustaceans 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 tropical reefs and sponges.[67] 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.[68]

Human eusociality

E. O. Wilson and Bert Hölldobler controversially[69] 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.[70]

See also
Collectivism Nesting instinct
Dominance hierarchy Prosocial behavior
Group cohesiveness Reciprocal altruism
Group selection Social behavior
Individualism Stigmergy
Interdependence

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