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. 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, 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.[3] Although presocial species are much more common than eusocial species, eusocial species have disproportionately large populations.[4]

Canis lupus pack surrounding Bison
Gray wolves hunting in a presocial pack encircle an American bison.

Presociality

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).[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]

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)[9] and Costa (2006) [10] give readable overviews.

Insect groups with species exhibiting subsociality include:

Dermaptera[11][12][13]

Orthoptera

Blattodea

Zoraptera[14]

Psocoptera[15]

Embioptera[16] [17].

Hemiptera[18][19]: many families, including Membracidae[20][21], Pentatomidae[22], Reduviidae[23][24][25], Tingidae[26]. Paternal care is known in some reduviids.[27]

Thysanoptera

Coleoptera [28]. Species and clades in multiple families: Erotylidae[29], Scarabaeidae[30] [31], Silvanidae,

Tenebrionidae[32][33],

Gyrinidae,

Passalidae[34][35][36] The biology of Odontotaenius disjunctus (Illiger) is summarized by Bibbs et al.[37],

Silphidae[38], Staphylinidae[39][40]

Curculionidae;

Scolytinae[41][42][43][44],

Platypodinae[45][43],

Other Curculionidae.

Chrysomelidae. In the leaf beetles, subsociality has been documented in two subfamilies, Cassidinae and Chrysomelinae.[46][47][48]. In Cassidinae (43 tribes, 325 genera, ~6000 species), maternal guarding of broods (eggs, larvae, pupae) is documented so far in two tribes (in the derived tortoise beetles clade[49]). In the Neotropical tribe Eugenysini, it appears in both Agenysa and Eugenysa[50]. In the tribe Mesomphaliini, subsocial species are documented in Acromis[51], Omaspides, and Paraselenis. In Chrysomelinae (150 genera, ~3000 species), maternal guarding of eggs and larvae is known in three tribes: Doryphorini: Doryphora and Platyphora; Gonioctenini: Gonioctena; and Phyllocharitini: Pterodunga mirabile in Australia[52]. See Chaboo et al. 2014 for a molecular-based phylogeny of Chrysomelinae and discussion of the origin of maternal care. Generally, reproductive and defense strategies in this subfamily are very diverse, with situations of ovoviviparity and viviparity and chemical defense[53].

Lepidoptera[54]

Hymenoptera

Solitary but social

Microcebus murinus -Artis Zoo, Amsterdam, Netherlands-8c
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.[55]

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.[3]

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,[56] as well as in some other invertebrates.)[3] 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
Giant honey bees cover the honeycomb of their nest.

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 incompertus), Hemiptera (bugs such as Pemphigus spyrothecae), and Thysanoptera (thrips) are described as eusocial. Eusocial species that lack this criterion of morphological caste differentiation are said to be primitively eusocial.[3]

Two potential examples of primitively eusocial mammals are the naked mole-rat and the Damaraland mole-rat (Heterocephalus glaber & Fukomys damarensis, respectively).[57] 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.[58]

Eusociality has arisen among some crustaceans that live in groups in a restricted area. Synalpheus regalis are parasitic shrimp that rely on fortress defense. They live in groups of closely related individuals, amidst tropical reefs and sponges.[59] 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.[60]

Human eusociality

E. O. Wilson and Bert Hölldobler controversially[61] 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.[62]

Comparison of sociality classifications

Degree of sociality Parental investment Cohabitation of all
adults and young
Cooperative care
of young
Reproductive division of labour Caste system Overlapping adult generations
Solitary
Subsocial Yes No No No No No
Solitary but social Yes Sometimes No No No No
Parasocial
Communal Yes Yes No No No No
Quasisocial Yes Yes Yes No No No
Semisocial Yes Yes Yes Yes Yes No
Eusocial Yes Yes Yes Yes Yes Yes

See also

References

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