Sociality

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Gray wolves hunting in a presocial pack encircle an American bison.

Sociality is the degree to which individuals in an animal population tend to associate in social groups 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. Parental investment is any expenditure of resources (time, 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.

Solitary animals, such as the jaguar, don't 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.^[3] Although presocial species are much more common than eusocial species, eusocial species have disproportionately large populations.^[4]

Presociality

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).^[5][6] 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.^[7]

An example of a species that exhibits presociality is the wasp species Philanthus gibbosus. P. gibbosus exhibits presociality in that it lives communally only for a very short amount of time while establishing a new burrow and at other certain phases of the nesting cycle. Apart from those specific points in the nesting cycle, P. gibbosus has been observed to live completely solitarily.^[8]

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.

Solitary but social

The mouse lemur is a nocturnal, solitary-but-social lemur native to Madagascar.

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. Some examples of solitary-but-social species are mouse lemurs, lorises, and orangutans.^[9]

Parasociality

Sociobiologists place communal, quasisocial, and semisocial animals into a meta-class: the parasocial. What parasocial taxa have in common is that they socialize in a single, cooperative dwelling.^[3]

Here is how communal, quasisocial, and semisocial taxa differ: In a communal group, adults cohabit in a single nest site, but they each care for their own young. Quasisocial animals, however, additionally share the responsibilities of brood care.

Quasisocial behaviour has been observed in some Hymenoptera and spider taxa,^[10] as well as in some other invertebrates.^[3]

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

Giant honey bees cover the honeycomb of their hive.

Eusocial societies have overlapping adult generations, cooperative care of young, and division of labor. However, in some animal species the division of labour is more flexible than in others. An individual's duty is dictated by caste, but one's caste may change.

For example, a queen bee can become a worker bee, and a worker bee can become a queen bee.^[3][11] The individual has a social role, but that role is not codified in the shape of the organism (its morphology). However, when organisms in a species are born with physical characteristics specific to a caste, and that caste never changes in the life of the organism, this exemplifies the highest acknowledged degree of sociality. Only two orders in the animal kingdom include highly eusocial taxa: Hymenoptera (ants, bees, sawflies, and wasps) and the infraorder isoptera (termites). Eusocial species that lack this criterion of morphological caste differentiation are said to be primitively eusocial.^[3]

Mammalian examples include the naked mole-rat and the Damaraland mole-rat (Heterocephalus glaber & Fukomys damarensis, respectively),^[12] two species of vertebrates that are diploid and highly inbred. Usually living in harsh or limiting environments, these mole rats aid in raising siblings and relatives born to a single reproductive queen. However, this classification is controversial owing to disputed definitions of 'eusociality.' 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.^[13] Thus, it is uncertain whether mole rats classify as true eusocial organisms, since their social behavior depends largely on their resources and environment.

Eusociality has also arisen among some crustaceans that live in groups in a restricted area. Synalpheus regalis, parasitic shrimp that rely on fortress defense and live in groups of closely related individuals in tropical reefs and sponges,^[14] live eusocially with a single breeding female and a large number of male defenders, 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.^[15]

Comparison of sociality classifications

See also

References