Sociality

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Gray wolves hunting in a presocial pack encircle an American bison. Canis lupus pack surrounding Bison.jpg
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 (gregariousness) and form cooperative societies.

Contents

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.

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]

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 bold italics on a phylogenetic tree of the Neoptera (note that many non-subsocial groups are omitted): [12]

Neoptera
Idioprothoraca

Embioptera (webspinners) [13] [14]

Rhipineoptera
Dictyoptera

Blattodea (cockroaches, inc. eusocial termites) [15]

Mantodea (mantises)

Orthoptera (grasshoppers, crickets) [16]

Dermaptera (earwigs) [17] [18] [19]

Eumetabola
Parametabola

Zoraptera (angel insects) [20]

Paraneoptera
Condylognatha

Thysanoptera (thrips) [21]

Hemiptera  (bugs)

Membracidae (treehoppers, thorn bugs) [22] [23]

Pentatomidae (shield bugs) [24]

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

Tingidae (lace bugs) [28] [29]

many families [30] [31]

Psocoptera (bark lice) [32]

Endopterygota
Coleoptera [33]

Staphylinidae (rove beetles) [34] [35] [36] [37] [38] [39] [40]

Silphidae (carrion beetles) [41]

Passalidae (bessbugs) [42] [43] [44] [45]

Scarabaeidae (scarabs) [46] [47]

Tenebrionidae (leaf/flower beetles) [48] [49]

Erotylidae (pleasing fungus beetles) [50]

Chrysomelidae (leaf beetles) [51] [52] [53] [54] [55] [56] [57] [58] [59]

Neuropteroidea

Raphidioptera (snakeflies)

Neuroptera (lacewings, alderflies, and allies)

Antliophora (true flies, scorpionflies, fleas)

Trichoptera (caddisflies)

Lepidoptera (butterflies and moths) [60]

Hymenoptera (sawflies, wasps, ants, bees) [61] (apart from eusocial species)

Solitary but social

The mouse lemur is a nocturnal, solitary-but-social lemur native to Madagascar. Microcebus murinus -Artis Zoo, Amsterdam, Netherlands-8c.jpg
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. However, this is opposite among cassowaries, for example. 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]

Some individual cetaceans 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, 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

Giant honey bees cover the honeycomb of their nest. Natural Beehive and Honeycombs.jpg
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. [4]

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. 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

Related Research Articles

<span class="mw-page-title-main">Beetle</span> Order of insects

Beetles are insects that form the order Coleoptera, in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 described species, is the largest of all orders, constituting almost 40% of described insects and 25% of all known animal species; new species are discovered frequently, with estimates suggesting that there are between 0.9 and 2.1 million total species. Found in almost every habitat except the sea and the polar regions, they interact with their ecosystems in several ways: beetles often feed on plants and fungi, break down animal and plant debris, and eat other invertebrates. Some species are serious agricultural pests, such as the Colorado potato beetle, while others such as Coccinellidae eat aphids, scale insects, thrips, and other plant-sucking insects that damage crops.

<span class="mw-page-title-main">Silphidae</span> Family of beetles

Silphidae is a family of beetles that are known commonly as large carrion beetles, carrion beetles or burying beetles. There are two subfamilies: Silphinae and Nicrophorinae. Nicrophorines are sometimes known as sexton beetles. The number of species is relatively small at around two hundred. They are more diverse in the temperate region although a few tropical endemics are known. Both subfamilies feed on decaying organic matter such as dead animals. The subfamilies differ in which uses parental care and which types of carcasses they prefer. Silphidae are considered to be of importance to forensic entomologists because when they are found on a decaying body they are used to help estimate a post-mortem interval.

<span class="mw-page-title-main">Vespidae</span> Family of insects

The Vespidae are a large, diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps and many solitary wasps. Each social wasp colony includes a queen and a number of female workers with varying degrees of sterility relative to the queen. In temperate social species, colonies usually last only one year, dying at the onset of winter. New queens and males (drones) are produced towards the end of the summer, and after mating, the queens hibernate over winter in cracks or other sheltered locations. The nests of most species are constructed out of mud, but polistines and vespines use plant fibers, chewed to form a sort of paper. Many species are pollen vectors contributing to the pollination of several plants, being potential or even effective pollinators, while others are notable predators of pest insect species, and a few species are invasive pests.

A trophic egg is an egg whose function is not reproduction but nutrition; in essence, the trophic egg serves as food for offspring hatched from viable eggs. In most species that produce them, a trophic egg is usually an unfertilised egg. The production of trophic eggs has been observed in a highly diverse range of species, including fish, amphibians, spiders and insects. The function is not limited to any particular level of parental care, but occurs in some sub-social species of insects, the spider A. ferox, and a few other species like the frogs Leptodactylus fallax and Oophaga, and the catfish Bagrus meridionalis.

<span class="mw-page-title-main">Social caterpillars</span> Behaviors of caterpillars in society

The collective behaviors of social caterpillars falls into five general categories: collective and cooperative foraging, group defense against predators and parasitoids, shelter building, thermoregulation and substrate silking to enhance steadfastness.

<i>Neochlamisus</i> Genus of beetles

Neochlamisus is a genus of leaf beetles in the tribe Fulcidacini. They are members of the case-bearing leaf beetle group, the Camptosomata. Measuring 3–4 millimeters in length as adults, they are cryptic, superficially resembling caterpillar frass. Seventeen, sometimes 18 species are presently accepted in this genus, all of them occurring in North America.

<span class="mw-page-title-main">Cassidinae</span> Subfamily of beetles

The Cassidinae are a subfamily of the leaf beetles, or Chrysomelidae. The antennae arise close to each other and some members have the pronotal and elytral edges extended to the side and covering the legs so as to give them the common name of tortoise beetles. Some members, such as in the tribe Hispini, are notable for the spiny outgrowths to the pronotum and elytra.

<span class="mw-page-title-main">Parental care</span>

Parental care is a behavioural and evolutionary strategy adopted by some animals, involving a parental investment being made to the evolutionary fitness of offspring. Patterns of parental care are widespread and highly diverse across the animal kingdom. There is great variation in different animal groups in terms of how parents care for offspring, and the amount of resources invested by parents. For example, there may be considerable variation in the amount of care invested by each sex, where females may invest more in some species, males invest more in others, or investment may be shared equally. Numerous hypotheses have been proposed to describe this variation and patterns in parental care that exist between the sexes, as well as among species.

<span class="mw-page-title-main">Mass provisioning</span>

Mass provisioning is a form of parental investment in which an adult insect, most commonly a hymenopteran such as a bee or wasp, stocks all the food for each of her offspring in a small chamber before she lays the egg. This behavior is common in both solitary and eusocial bees, though essentially absent in eusocial wasps.

<span class="mw-page-title-main">Eusociality</span> Highest level of animal sociality a species can attain

Eusociality, the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care, overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups. The division of labor creates specialized behavioral groups within an animal society which are sometimes referred to as 'castes'. Eusociality is distinguished from all other social systems because individuals of at least one caste usually lose the ability to perform at least one behavior characteristic of individuals in another caste. Eusocial colonies can be viewed as superorganisms.

<i>Gargaphia solani</i> Species of true bug

Gargaphia solani is a subsocial species of lace bug commonly known as the eggplant lace bug. The species was described by Heidemann in 1914 after it aroused attention a year earlier in the United States as an eggplant pest around Norfolk, Virginia. Fink found that the species became an agricultural pest when eggplant is planted on a large scale.

<i>Austroplatypus incompertus</i> Species of beetle

Austroplatypus incompertus is a species of ambrosia beetle belonging to the true weevil family, native to Australia, with a verified distribution in New South Wales and Victoria. It forms colonies in the heartwood of Eucalyptus trees and is the first beetle to be recognized as a eusocial insect. Austroplatypus incompertus is considered eusocial because groups contain a single fertilized female that is protected and taken care of by a small number of unfertilized females that also do much of the work. The species likely passed on cultivated fungi to other weevils.

<span class="mw-page-title-main">Evolution of eusociality</span> Origins of cooperative brood care

Eusociality evolved repeatedly in different orders of animals, notably termites and the Hymenoptera. This 'true sociality' in animals, in which sterile individuals work to further the reproductive success of others, is found in termites, ambrosia beetles, gall-dwelling aphids, thrips, marine sponge-dwelling shrimp, naked mole-rats, and many genera in the insect order Hymenoptera. The fact that eusociality has evolved so often in the Hymenoptera, but remains rare throughout the rest of the animal kingdom, has made its evolution a topic of debate among evolutionary biologists. Eusocial organisms at first appear to behave in stark contrast with simple interpretations of Darwinian evolution: passing on one's genes to the next generation, or fitness, is a central idea in evolutionary biology.

<i>Appasus</i> Genus of true bugs

Appasus is a genus of giant water bugs found in freshwater habitats in Asia and Africa.

<i>Abedus</i> Genus of true bugs

Abedus is a genus of giant water bugs found in freshwater habitats in southern United States, Mexico and Central America. Sometimes called ferocious water bugs, these brown insects typically are between 2.3 and 4 cm (0.9–1.6 in) long, although A. immaculatus only is about 1.3–1.4 cm (0.51–0.55 in), making it the smallest North American belostomatid and possibly worthy of separation in its own genus. Otherwise the different Abedus species are very similar and can often only be separated with a microscope. They will bite in self-defense, which is painful but not dangerous.

In biology, paternal care is parental investment provided by a male to his own offspring. It is a complex social behaviour in vertebrates associated with animal mating systems, life history traits, and ecology. Paternal care may be provided in concert with the mother or, more rarely, by the male alone.

Kladothrips is a genus of Australian gall thrips. It is notable for including some of the few organisms outside of Hymenoptera that exhibit eusociality.

<span class="mw-page-title-main">Social immunity</span> Antiparasite defence mounted for the benefit of individuals other than the actor

Social immunity is any antiparasite defence mounted for the benefit of individuals other than the actor. For parasites, the frequent contact, high population density and low genetic variability makes social groups of organisms a promising target for infection: this has driven the evolution of collective and cooperative anti-parasite mechanisms that both prevent the establishment of and reduce the damage of diseases among group members. Social immune mechanisms range from the prophylactic, such as burying beetles smearing their carcasses with antimicrobials or termites fumigating their nests with naphthalene, to the active defenses seen in the imprisoning of parasitic beetles by honeybees or by the miniature 'hitchhiking' leafcutter ants which travel on larger worker's leaves to fight off parasitoid flies. Whilst many specific social immune mechanisms had been studied in relative isolation, it was not until Sylvia Cremer et al.'s 2007 paper "Social Immunity" that the topic was seriously considered. Empirical and theoretical work in social immunity continues to reveal not only new mechanisms of protection but also implications for understanding of the evolution of group living and polyandry.

<i>Proseicela</i> Genus of beetles

Proseicela is a genus of leaf beetles.

<i>Doryphora</i> (beetle) Genus of leaf beetles

Doryphora is a genus of leaf beetles in the family Chrysomelidae. It includes nine species from Central and South America.

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