727:, has evolved eusociality involving a colony creating a class of sterile soldiers. One fluke invades a host and establishes a colony of dozens to thousands of clones that work together to take it over. Since rival trematode species can invade and replace the colony, it is protected by a specialized caste of sterile soldier trematodes. Soldiers are smaller, more mobile, and develop along a different pathway than sexually mature reproductives. One difference is that a soldier's mouthparts (pharynx) is five times as big as those of the reproductives. They make up nearly a quarter of the volume of the soldier. These soldiers do not have a germinal mass, never metamorphose to be reproductive, and are, therefore, obligately sterile. Soldiers are readily distinguished from the immature and mature reproductive worms. Soldiers are more aggressive than reproductives, attacking heterospecific trematodes that infect their host
745:
1138:
1730:(actual or potential queens) can cause size dimorphisms between different castes, as size is strongly influenced by the season during which the individual is reared. In many wasps, worker caste is determined by a temporal pattern in which workers precede non-workers of the same generation. In some cases, for example in bumblebees, queen control weakens late in the season, and the ovaries of workers develop. The queen attempts to maintain her dominance by aggressive behavior and by eating worker-laid eggs; her aggression is often directed towards the worker with the greatest ovarian development.
999:
1501:
to what W. D. Hamilton first termed "supersisters", more closely related to their sisters than they would be to their own offspring. Even though workers often do not reproduce, they can pass on more of their genes by helping to raise their sisters than by having their own offspring (each of which would only have 50% of their genes). This unusual situation, where females may have greater fitness when they help rear sisters rather than producing offspring, is often invoked to explain the multiple independent evolutions of eusociality (at least nine separate times) within the
Hymenoptera.
1465:
280:, the mole-rats. Further research distinguished another possibly important criterion for eusociality, "the point of no return". This is characterized by having individuals fixed into one behavioral group, usually before reproductive maturity. This prevents them from transitioning between behavioral groups, and creates a society with individuals truly dependent on each other for survival and reproductive success. For many insects, this irreversibility has changed the anatomy of the worker caste, which is sterile and provides support for the reproductive caste.
520:
1637:
1078:
1583:
bees was followed by at least one reversal to solitarity, giving a total of at least nine reversals. In a few species, solitary and eusocial colonies appear simultaneously in the same population, and different populations of the same species may be fully solitary or eusocial. This suggests that eusociality is costly to maintain, and can only persist when ecological variables favor it. Disadvantages of eusociality include the cost of investing in non-reproductive offspring, and an increased risk of disease.
1162:
1298:
575:
1518:
association between haplodiploidy and eusociality is below statistical significance. Haplodiploidy is thus neither necessary nor sufficient for eusociality to emerge. Relatedness does still play a part, as monogamy (queens mating singly) is the ancestral state for all eusocial species so far investigated. If kin selection is an important force driving the evolution of eusociality, monogamy should be the ancestral state, because it maximizes the relatedness of colony members.
327:
174:
33:
6890:
1240:
1035:
374:, dominant females perform tasks such as building new cells and ovipositing, while subordinate females tend to perform tasks like feeding the larvae and foraging. The task differentiation between castes can be seen in the fact that subordinates complete 81.4% of the total foraging activity, while dominants only complete 18.6% of the total foraging. Eusocial species with a sterile caste are sometimes called hypersocial.
1206:
6675:
6389:
541:) make up another large portion of highly advanced eusocial animals. The colony is differentiated into various castes: the queen and king are the sole reproducing individuals; workers forage and maintain food and resources; and soldiers defend the colony against ant attacks. The latter two castes, which are sterile and perform highly specialized, complex social behaviors, are derived from different stages of
6699:
249:
1738:, which is secreted by workers, but normally they are switched over to a diet of pollen and honey as they mature; if their diet is exclusively royal jelly, they grow larger than normal and differentiate into queens. This jelly contains a specific protein, royalactin, which increases body size, promotes ovary development, and shortens the developmental time period. The differential expression in
1633:, consistent with their roles as sex attractants during the mating flight. Once a queen is mated and begins laying eggs, she starts producing the full blend of compounds. In several ant species, reproductive activity is associated with pheromone production by queens. Mated egg-laying queens are attractive to workers, whereas young winged virgin queens elicit little or no response.
5720:
1547:, social phenotype has been predicted by altitude and micro-habitat composition, with social nests found in warmer, sunnier sites, and solitary nests found in adjacent, cooler, shaded locations. Facultatively social bee species, however, which comprise the majority of social bee diversity, have their lowest diversity in the tropics, being largely limited to temperate regions.
1270:
784:, mole rats sometimes outbreed and establish new colonies when resources are sufficient. Most of the individuals cooperatively care for the brood of a single reproductive female (the queen) to which they are most likely related. Thus, it is uncertain whether mole rats are truly eusocial, since their social behavior depends largely on their resources and environment.
878:
6687:
1539:, where eusociality has been gained and lost multiple times, are correlated with periods of climatic warming. Social behavior in facultative social bees is often reliably predicted by ecological conditions, and switches in behavioral type have been experimentally induced by translocating offspring of solitary or social populations to warm and cool climates. In
1655:
development in new winged females, called female sexuals. These chemicals inhibit workers from rearing male and female sexuals, suppress egg production in other queens of multiple queen colonies, and cause workers to execute excess queens. These pheromones maintain the eusocial phenotype, with one queen supported by sterile workers and sexually active males (
1389:
505:, a halictid bee from Central America, nests may be headed by more than one female; such nests have more cells, and the number of active cells per female is correlated with the number of females in the nest, implying that having more females leads to more efficient building and provisioning of cells. In similar species with only one queen, such as
451:
1618:, which control workers. Mandibular gland extracts inhibit workers from constructing queen cells, which can delay the hormonally based behavioral development of workers and suppress their ovarian development. Both behavioral effects mediated by the nervous system often leading to recognition of queens (
566:) to be recognized as eusocial. This species forms colonies in which a single female is fertilized, and is protected by many unfertilized females, which serve as workers excavating tunnels in trees. This species has cooperative brood care, in which individuals care for juveniles that are not their own.
1733:
In highly eusocial wasps (where castes are morphologically dissimilar), both the quantity and quality of food are important for caste differentiation. Recent studies in wasps suggest that differential larval nourishment may be the environmental trigger for larval divergence into workers or gynes. All
1603:
play an important role in the physiological mechanisms of eusociality. Enzymes involved in the production and perception of pheromones were important for the emergence of eusociality within both termites and hymenopterans. The best-studied queen pheromone system in social insects is that of the honey
1586:
All reversals to solitarity have occurred among primitively eusocial groups; none have followed the emergence of advanced eusociality. The "point of no return" hypothesis posits that the morphological differentiation of reproductive and non-reproductive castes prevents highly eusocial species such as
1517:
Further, not all eusocial species are haplodiploid: termites, some snapping shrimps, and mole rats are not. Conversely, many non-eusocial bees are haplodiploid, and among eusocial species many queens mate with multiple males, resulting in a hive of half-sisters that share only 25% of their genes. The
1488:
theory, organisms can gain fitness by increasing the reproductive output of other individuals that share their genes, especially their close relatives. Natural selection favors individuals to help their relatives when the cost of helping is less than the benefit gained by their relative multiplied by
1725:
In primitively eusocial bees (where castes are morphologically similar and colonies are small and short-lived), queens frequently nudge their nest mates and then burrow back down into the nest. This draws workers into the lower part of the nest where they may respond to stimuli for cell construction
1500:
In haplodiploid species, females develop from fertilized eggs and males develop from unfertilized eggs. Because a male is haploid, his daughters share 100% of his genes and 50% of their mother's. Therefore, they share 75% of their genes with each other. This mechanism of sex determination gives rise
705:
The fortress defense hypothesis additionally points out that because sponges provide both food and shelter, there is an aggregation of relatives (because the shrimp do not have to disperse to find food), and much competition for those nesting sites. Being the target of attack promotes a good defense
1713:
wasps, monogamy is established soon after colony formation by physical dominance interactions among foundresses of the colony including biting, chasing, and food soliciting. Such interactions create a dominance hierarchy headed by larger, older individuals with the greatest ovarian development. The
234:) in order to investigate the different levels of animal sociality, all of which are different stages that a colony may pass through. Eusociality, which is the highest level of animal sociality a species can attain, specifically had three characteristics that distinguished it from the other levels:
202:
bees, on a scale of subsocial/solitary, colonial/communal, semisocial, and eusocial, where a colony is started by a single individual. Batra observed the cooperative behavior of the bees, males and females alike, as they took responsibility for at least one duty (i.e., burrowing, cell construction,
1582:
A reversal to solitarity is an evolutionary phenomenon in which descendants of a eusocial group evolve solitary behavior once again. Bees have been model organisms for the study of reversal to solitarity, because of the diversity of their social systems. Each of the four origins of eusociality in
701:
are the eight recorded species of parasitic shrimp that rely on fortress defense and live in groups of closely related individuals in tropical reefs and sponges. They live eusocially with a single breeding female, and a large number of male defenders armed with enlarged snapping claws. There is a
130:
take the roles of the sole reproducers, while soldiers and workers work together to create and maintain a living situation favorable for the brood. Queens produce multiple queen pheromones to create and maintain the eusocial state in their colonies; they may also eat eggs laid by other females or
1526:
Increased parasitism and predation rates are the primary ecological drivers of social organization. Group living affords colony members defense against enemies, specifically predators, parasites, and competitors, and allows them to gain advantage from superior foraging methods. The importance of
735:
soldiers do not attack conspecifics from other colonies. The soldiers are not evenly distributed throughout the host body. They are found in the highest numbers in the basal visceral mass, where competing trematodes tend to multiply during the early phase of infection. This strategic positioning
79:
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
1513:
notes that while females share 75% of genes with their sisters in haplodiploid populations, they only share 25% of their genes with their brothers. Accordingly, the average relatedness of an individual to their sibling is 50%. Therefore, helping behavior is only advantageous if it is biased to
1654:
includes both releaser and primer pheromones. A queen recognition (releaser) pheromone is stored in the poison sac along with three other compounds. These compounds elicit a behavioral response from workers. Several primer effects have also been demonstrated. Pheromones initiate reproductive
1569:
proposed in 2010 that since eusociality produces an extremely altruistic society, eusocial groups should out-reproduce their less cooperative competitors, eventually eliminating all non-eusocial groups from a species. Multilevel selection has been heavily criticized for its conflict with the
914:
species has both types. At the bottom of a colony, there are "nest" fronds that clasp the trunk of the tree supporting the fern, and drooping photosynthetic fronds. These are argued to be adapted to support the colony structurally, i.e. that the individuals in the colony are to some degree
478:
fill their abdomens with liquid food until they become immobile and hang from the ceilings of the underground nests, acting as food storage for the rest of the colony. Not all social insects have distinct morphological differences between castes. For example, in the
Neotropical social wasp
1659:). In queenless colonies, the lack of queen pheromones causes winged females to quickly shed their wings, develop ovaries and lay eggs. These virgin replacement queens assume the role of the queen and start to produce queen pheromones. Similarly, queen weaver ants
1431:
referred to the existence of sterile castes as the "one special difficulty, which at first appeared to me insuperable, and actually fatal to my theory". Darwin anticipated that a possible resolution to the paradox might lie in the close family relationship, which
909:
of differing size and shape, to collect and store water and nutrients for the colony to use. At the top of a colony, there are both pleated fan-shaped "nest" fronds that collect and hold water, and gutter-shaped "strap" fronds that channel water: no solitary
736:
allows them to effectively defend against invaders, similar to how soldier distribution patterns are seen in other animals with defensive castes. They "appear to be an obligately sterile physical caste, akin to that of the most advanced social insects".
463:
Reproductive specialization generally involves the production of sterile members of the species, which carry out specialized tasks to care for the reproductive members. Individuals may have behavior and morphology modified for group defense, including
1534:
Climate too appears to be a selective agent driving social complexity; across bee lineages and
Hymenoptera in general, higher forms of sociality are more likely to occur in tropical than temperate environments. Similarly, social transitions within
837:
cooperated to rear their children while other members of the same group hunted and foraged. Wilson and others argued that through cooperation and teamwork, ants and humans form superorganisms. Wilson's claims were vigorously rejected by critics of
956:
period (~150 million years ago). The other orders shown contain both eusocial and non-eusocial species, including many lineages where eusociality is inferred to be the ancestral state. Thus the number of independent evolutions of eusociality
780:. 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'. To
1440:
theory. After the gene-centered view of evolution was developed in the mid-1970s, non-reproductive individuals were seen as an extended phenotype of the genes, which are the primary beneficiaries of natural selection.
1421:, eusociality was seen as paradoxical: if adaptive evolution unfolds by differential reproduction of individual organisms, the evolution of individuals incapable of passing on their genes presents a challenge. In
1514:
helping sisters, which would drive the population to a 1:3 sex ratio of males to females. At this ratio, males, as the rarer sex, increase in reproductive value, reducing the benefit of female-biased investment.
1527:
ecology in the evolution of eusociality is supported by evidence such as experimentally induced reproductive division of labor, for example when normally solitary queens are forced together. Conversely, female
2833:
Wcislo, W. T.; Wille, A.; Orozco, E. (1993). "Nesting biology of tropical solitary and social sweat bees, Lasioglossum (Dialictus) figueresi Wcislo and L. (D.) aeneiventre (Friese) (Hymenoptera: Halictidae)".
811:). These show cooperative breeding and marked reproductive skews. In the dwarf mongoose, the breeding pair receives food priority and protection from subordinates and rarely has to defend against predators.
1678:. For a queen to dominate all the workers, usually numbering more than 3000 in a colony, she signals her dominance with pheromones. The workers regularly lick the queen while feeding her, and the air-borne
491:
workers. Between approximately 0β40 days old, the workers perform tasks within the nest such as provisioning cell broods, colony cleaning, and nectar reception and dehydration. Once older than 40 days,
262:
extended the terminology to include other social insects, such as ants, wasps, and termites. Originally, it was defined to include organisms (only invertebrates) that had the following three features:
1726:
and maintenance. Being nudged by the queen may help to inhibit ovarian development; in addition, the queen eats any eggs laid by workers. Furthermore, temporally discrete production of workers and
709:
Eusociality offers a competitive advantage in shrimp populations. Eusocial species are more abundant, occupy more of the habitat, and use more of the available resources than non-eusocial species.
1543:, females produce a single brood in cooler regions and two or more broods in warmer regions, so the former populations are solitary while the latter are social. In another species of sweat bees,
1742:
of larval genes and proteins (also differentially expressed during queen versus caste development in honey bees) indicates that regulatory mechanisms may operate very early in development.
2869:
Richards, Miriam H. (2000). "Evidence for geographic variation in colony social organization in an obligately social sweat bee, Lasioglossum malachurum Kirby (Hymenoptera; Halictidae)".
3776:
Chu, Carol; Buchman-Schmitt, Jennifer M.; Stanley, Ian H.; Hom, Melanie A.; Tucker, Raymond P.; Hagan, Christopher R.; Rogers, Megan L.; Podlogar, Matthew C.; Chiurliza, Bruno (2017).
545:
larvae produced by the reproductive caste. Some soldiers have jaws so enlarged (specialized for defense and attack) that they are unable to feed themselves and must be fed by workers.
4757:
Hughes, William O. H.; Benjamin P. Oldroyd; Madeleine
Beekman; Francis L. W. Ratnieks (2008-05-30). "Ancestral Monogamy Shows Kin Selection Is Key to the Evolution of Eusociality".
905:), may exhibit a primitive form of eusocial behavior amongst clones. The evidence for this is that individuals live in colonies, where they are structured in different ways, with
1699:. With low juvenile hormone, eggs do not mature. Similar inhibitory effects of lowering juvenile hormone were seen in halictine bees and polistine wasps, but not in honey bees.
1644:: three queen pheromones help to create and maintain the eusocial state of the colony. Loss of a primer pheromone triggers the development of replacement queens (dashed lines).
643:), but the gall-inhabiting behavior gives these species a defensible resource. They produce soldier castes for fortress defense and protection of the colony against predators,
1626:) are attributed to the same pheromones. These pheromones volatilize or are deactivated within thirty minutes, allowing workers to respond rapidly to the loss of their queen.
1803:
207:) within the colony. The cooperativeness was essential as the activity of one labor division greatly influenced the activity of another. Eusocial colonies can be viewed as
7232:
1836:
223:; castes fulfill a specific role that contributes to the functioning and survival of the whole colony, while being incapable of independent survival outside the colony.
1773:, asks whether Wilson's stated "wish" for humans to bring about "a permanent paradise for human beings" would mean "to be group-selected in factories in the style of
952:(non-eusocial groups not shown). All species of termites are eusocial, and it is believed that they were the first eusocial animals to evolve, sometime in the upper
2678:
Wongvilas, S.; Deowanish, S.; Lim, J.; Xie, V. R. D.; Griffith, O. W.; Oldroyd, B. P. (2010). "Interspecific and conspecific colony mergers in the dwarf honey bees
1902:
6323:
485:, caste ranks are determined by social displays in the developing brood. These castes are sometimes further specialized in their behavior based on age, as in
2518:
Richards, Miriam H. (2019). "Social trait definitions influence evolutionary inferences: a phylogenetic approach to improving social terminology for bees".
1718:
when queens are present, for a variety of reasons: colonies tend to be small enough that queens can effectively dominate workers; queens practice selective
1480:
females. It has been suggested that this organisation favours eusociality, but haplodiploidy is neither necessary nor sufficient for eusociality to emerge.
6328:
6921:
1707:
A variety of other mechanisms give queens of different species of social insects a measure of reproductive control over their nest mates. In many
5526:
2956:
7101:
6425:
5124:
Zara, Fernando; Balestieri, Jose (2000). "Behavioural
Catalogue of Polistes versicolor Olivier (Vespidae: Polistinae) Post-emergent Colonies".
2499:
Zara, Fernando; Balestieri, Jose (2000). "Behavioural
Catalogue of Polistes versicolor Olivier (Vespidae: Polistinae) Post-emergent Colonies".
5548:
Fromm, Harold (2013). "Review of
Groping for Groups, by Edward O. Wilson, Jonathan Haidt, Steven Mithen, Steven Pinker, and Richard Dawkins".
6840:
5611:
4022:"Inclusive fitness and differential productivity across the life course determine intergenerational transfers in a small-scale human society"
3977:
Hawkes, Kristen; Coxworth, James E. (2013). "Grandmothers and the evolution of human longevity: a review of findings and future directions".
3605:
3488:
1808:
4349:
Thorne, B.L.; Grimaldi, D.A.; Krishna, K. (2001) . "Early fossil history of the termites". In Abe, T.; Bignell, D.E; Higashi, M. (eds.).
6478:
5361:
Feyereisen, R.; Tobe, S. (1981). "A rapid partition assay for routine analysis of juvenile hormone released by insect corpora allata".
6850:
6737:
5974:
5584:
4397:
3422:
441:(red dwarf honey bee). Pheromones are sometimes used in these castes to assist with foraging. Workers of the Australian stingless bee
309:
3934:
VanderLaan, Doug P.; Ren, Zhiyuan; Vasey, Paul L. (2013). "Male androphilia in the ancestral environment. An ethnological analysis".
1685:
The mode of action of inhibitory pheromones which prevent the development of eggs in workers has been demonstrated in the bumble bee
230:
further expanded Batra's classification with his comparative study of social behavior in bees. He observed multiple species of bees (
7043:
2057:
3293:"Biodiversity, host specificity, and dominance by eusocial species among sponge-dwelling alpheid shrimp on the Belize Barrier Reef"
318:
Eusociality has evolved multiple times in different insect orders, including hymenopterans, termites, thrips, aphids, and beetles.
3248:
Hultgren, K.M.; Duffy, J. E. (2012). "Phylogenetic community ecology and the role of social dominance in sponge-dwelling shrimp".
647:, and competitors. In these groups, eusociality is produced by both high relatedness and by living in a restricted, shared area.
6940:
6612:
5921:
2783:
West-Eberhard, M. J. (1982). "The Nature and
Evolution of Swarming In Tropical Social Wasps (Vespidae, Polistinae, Polybini)".
1813:
1418:
998:
744:
459:
honeypot ants, showing the repletes or plerergates, their abdomens swollen to store honey (top), with ordinary workers (bottom)
1561:
Once pre-adaptations such as group formation, nest building, high cost of dispersal, and morphological variation are present,
6032:
4427:
1798:
2721:
Bartareau, T. (1996). "Foraging
Behaviour of Trigona Carbonaria (Hymenoptera: Apidae) at Multiple-Choice Feeding Stations".
429:
that alter the behavior of specific castes in the colony. These pheromones may act across different species, as observed in
3112:
Duffy, J. Emmett; Morrison, Cheryl L.; Rios, Ruben (2000). "Multiple origins of eusociality among sponge-dwelling shrimps (
5149:
4849:
Molteno, A. J.; Bennett, N. C. (2002). "Rainfall, dispersal and reproductive inhibition in eusocial
Damaraland mole-rats (
3163:"On the frequency of eusociality in snapping shrimps (Decapoda: Alpheidae), with description of a second eusocial species"
2798:
van Veen, J. W.; Sommeijer, M. J.; Meeuwsen, F. (November 1997). "Behaviour of drones in Melipona (Apidae, Meliponinae)".
255:, here collaborating to pull nest leaves together, can be considered eusocial, as they have a permanent division of labor.
5403:"Differential gene expression and protein abundance evince ontogenetic bias toward castes in a primitively eusocial wasp"
5143:
Harrison, Mark C.; Jongepier, Evelien; Robertson, Hugh M.; Arning, Nicolas; Bitard-Feildel, Tristan; et al. (2018).
706:
system (soldier caste); soldiers promote the fitness of the whole nest by ensuring safety and reproduction of the queen.
6811:
6762:
6366:
6062:
3735:
3292:
3162:
869:. These species are not eusocial: they do not have castes, and helpers reproduce on their own if given the opportunity.
3374:"The physical soldier caste of an invasive, human-infecting flatworm is morphologically extreme and obligately sterile"
1137:
7253:
7058:
7011:
6703:
6144:
5687:
5289:
3850:
2871:
1988:
4161:"With a little help from her friends (and family) part I: the ecology and evolution of non-maternal care in mammals"
3696:
Kesebir, Selin (2012). "The Superorganism Account of Human Sociality: How and When Human Groups Are Like Beehives".
1464:
849:
Though controversial, it has been suggested that male homosexuality and female menopause could have evolved through
519:
6767:
6620:
6513:
6418:
6345:
6237:
6094:
5839:
5667:
4354:
4217:
3778:"The interpersonal theory of suicide: A systematic review and meta-analysis of a decade of cross-national research"
1200:
861:", with juveniles and sexually mature adolescents helping their parents raise subsequent broods, as in some birds,
554:
313:
3831:(24 May 2012). "The Descent of Edward Wilson. Book review of 'The Social Conquest of Earth' by Edward O. Wilson".
2983:
6945:
6916:
6816:
6089:
5604:
2913:
1930:
5316:
Carpenter, J.M (1987). "Phylogenetic relationships and classification of the Vespinae (Hymenoptera: Vespidae)".
4967:"Behavioral and genetic mechanisms of social evolution: insights from incipiently and facultatively social bees"
3471:
O'Riain, M.J.; Faulkes, C. G. (2008). "African Mole-Rats: Eusociality, Relatedness and Ecological Constraints".
2561:
Peters, Ralph S.; Krogmann, Lars; Mayer, Christoph; Donath, Alexander; Gunkel, Simon; et al. (April 2017).
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6571:
6535:
6359:
6042:
5634:
2049:
1423:
928:
673:
501:
1077:
6691:
2981:
Aoki, S.; Imai, M. (2005). "Factors affecting the proportion of sterile soldiers in growing aphid colonies".
7258:
7177:
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6498:
6468:
6458:
6017:
5897:
5692:
4389:
3340:
Richards, Miriam H. (10 September 2024). "Social evolution and reproductive castes in trematode parasites".
1829:
1823:
1750:
1636:
507:
1629:
The levels of two of the aliphatic compounds increase rapidly in virgin queens within the first week after
186:
7197:
6857:
6730:
6176:
6139:
6129:
6074:
6022:
5967:
1565:
has been suggested as a driver of the transition to advanced eusociality. M. A. Nowak, C. E. Tarnita, and
1383:
897:
883:
752:
burrow with soldiers, workers, and queen, a social structure similar to the castes of the eusocial insects
443:
220:
32:
7263:
7006:
6911:
6679:
6473:
6411:
5318:
4497:"A unified model of Hymenopteran preadaptations that trigger the evolutionary transition to eusociality"
3854:
1722:; the flow of nutrients favors queen over workers; and queens rapidly lay eggs in new or vacated cells.
1161:
613:
487:
397:) are eusocial. Some major lineages of wasps are mostly or entirely eusocial, including the subfamilies
332:
5221:
Vargo, E. (1999). "Reproductive development and ontogeny or queen pheromone production in the fire ant
2348:
Stern, D. L. (1994). "A phylogenetic analysis of soldier evolution in the aphid family Hormaphididae".
4212:
1924:
Opachaloemphan, Comzit; Yan, Hua; Leibholz, Alexandra; Desplan, Claude; Reinberg, Danny (2018-11-23).
1297:
635:), are described as eusocial. These species have very high relatedness among individuals due to their
7217:
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6448:
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6282:
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5158:
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5021:
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4672:
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4508:
4461:
4294:"On the selective advantage of coloniality in staghorn ferns (Platycerium bifurcatum, Polypodiaceae)"
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workers, and sometimes also soldiers that perform specialized tasks. In the well-studied social wasp
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6503:
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5937:
5647:
3201:"Kin structure, ecology and the evolution of social organization in shrimp: a comparative analysis"
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858:
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227:
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single shared living space for the colony members, and the non-breeding members act to defend it.
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6520:
6287:
6079:
6057:
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5242:
4947:
4884:"Evo-Devo and the evolution of social behavior: Brain gene expression analyses in social insects"
4855:
4820:
4792:
4637:
4242:
4190:
4002:
3959:
3537:
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3322:
3143:
3094:
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2435:
2381:
2223:
1650:
916:
843:
723:
667:
431:
149:
6550:
5910:
2146:
447:, for instance, mark food sources with a pheromone, helping their nest mates to find the food.
326:
161:
have evolved a weak form of eusociality. It has been suggested that the colonial and epiphytic
7172:
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2373:
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2215:
2207:
2182:
2124:
2053:
1965:
1947:
1769:
1687:
1485:
1455:
1437:
1239:
949:
4599:
80:
behaviors characteristic of individuals in another caste. Eusocial colonies can be viewed as
7053:
7001:
6984:
6906:
6898:
6657:
6646:
6508:
6352:
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6164:
5997:
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2807:
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2647:
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2527:
2427:
2365:
2323:
2315:
2271:
2261:
2199:
2114:
2104:
2078:
2037:
1997:
1955:
1939:
1877:
1793:
1763:, in which most individuals are subject to reproductive repression. Harold Fromm, reviewing
1696:
1674:
830:
481:
212:
3450:
3184:
Duffy, J. E. (2003). "The ecology and evolution of eusociality in sponge-dwelling shrimp".
944:
Eusociality is a rare but widespread phenomenon in species in at least seven orders in the
526:
live in large nests, with queen, king, soldier (red heads), and worker (pale heads) castes.
7030:
6869:
6754:
6746:
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6488:
6483:
6307:
6247:
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6217:
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5884:
5707:
5672:
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5652:
4381:
3828:
2567:
1779:
1714:
rank of subordinates is correlated with the degree of ovarian development. Workers do not
1562:
1556:
1034:
961:) is not known. The major eusocial groups are shown in boldface in the phylogenetic tree.
839:
820:
640:
559:
173:
4260:
511:
in Europe, the degree of eusociality depends on the clime in which the species is found.
435:(black dwarf honey bee), where worker bees responded to queen pheromone from the related
5477:
5420:
5331:
5162:
5084:
5025:
4926:
Yanega, D. (1993). "Environmental influences on male production and social structure in
4772:
4676:
4568:
4512:
4465:
4120:"Environmental Uncertainty and the Global Biogeography of Cooperative Breeding in Birds"
3665:"Clash of the Titans. Book review of 'The Social Conquest of Earth' by Edward O. Wilson"
3517:
3398:
3373:
3308:
3261:
3039:
2996:
2926:
2633:
2423:
2361:
2195:
2100:
2001:
1943:
833:
called humans eusocial apes, arguing for similarities to ants, and observing that early
7212:
7146:
7123:
6957:
6952:
6935:
6928:
6821:
6586:
6272:
6227:
6222:
6104:
6012:
5781:
5439:
5402:
5339:
5179:
5144:
5101:
5066:
5042:
5007:
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4552:
4529:
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4449:
4422:
4326:
4293:
4095:
4070:
4046:
4021:
3911:
3876:
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3777:
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3556:
3225:
3200:
3131:
3082:
2652:
2615:
2151:
2119:
2082:
1960:
1925:
1865:
1692:
1606:
1510:
1428:
1205:
804:
761:
749:
644:
136:
5462:
Kamakura, Masaki (May 2011). "Royalactin induces queen differentiation in honeybees".
2328:
2303:
2276:
2249:
7247:
7086:
6828:
6799:
6777:
6601:
6596:
6493:
6277:
6267:
6262:
6257:
6186:
6119:
6027:
5677:
5662:
5374:
5287:
Fletcher, D.; Ross, K. (1985). "Regulation of Reproduction in Eusocial Hymenoptera".
5238:
4473:
4020:
Hooper, Paul L.; Gurven, Michael; Winking, Jeffrey; Kaplan, Hillard S. (2015-03-22).
3846:
3317:
3269:
2960:
2547:
2458:
2227:
1898:
1774:
1571:
1459:
1433:
902:
862:
850:
474:
455:
208:
195:
177:
162:
81:
48:
5401:
Hunt, J.; Wolschin, F.; Henshaw, M.; Newman, T.; Toth, A.; Amdam, G. (17 May 2010).
5246:
4951:
4796:
4246:
4194:
4119:
4006:
3963:
3446:
3326:
2819:
2439:
2385:
2319:
2042:
1669:
glands that produce pheromones which prevent workers from laying reproductive eggs.
336:
ants. The young queens are black, winged, and much larger than the wingless workers.
7167:
7141:
7131:
7108:
6989:
6606:
6576:
6542:
6434:
6302:
6252:
6212:
6181:
6134:
5915:
5807:
5639:
5511:
5347:
4812:"The emergence of reproductive division of labor in forced queen groups of the ant
4641:
3541:
3147:
3098:
3055:
3012:
2855:
2707:
1760:
1566:
1494:
1493:. W. D. Hamilton suggested in 1964 that eusociality could evolve more easily among
1469:
1124:
1114:
854:
632:
469:
259:
154:
124:
76:
5302:
4309:
4230:
4177:
4160:
3893:
2911:
Adams, E. S. (1987). "Territory size and population limits in mangrove termites".
248:
5429:
4213:"Group size and the suppression of subordinate reproduction in Kalahari meerkats"
3480:
2642:
7136:
6977:
6833:
6651:
6242:
6207:
5813:
5749:
5734:
5702:
1903:"Nests and Social Behavior of Halictine bees of India (Hymenoptera: Halictidae)"
1735:
1656:
1531:
undergo hormonal changes that promote dispersal after periods of high rainfall.
1218:
1131:
661:
627:
542:
437:
386:
365:
361:
341:
252:
204:
104:
3877:"Kin and multilevel selection in social evolution: a never-ending controversy?"
3754:
3624:
2531:
6994:
6972:
6641:
6555:
5868:
5833:
5799:
5793:
5754:
5719:
5682:
5170:
4983:
4966:
4868:
4725:
4136:
4086:
3947:
3504:
O' Riain, M.; et al. (1996). "A Dispersive Morph in the Naked Mole-Rat".
3004:
2699:
2581:
2562:
2472:
Cervo, Rita (2006). "Polistes wasps and their social parasites: an overview".
1755:
1600:
1536:
1269:
1193:
1104:
1028:
1011:
757:
656:
563:
426:
425:(orchid bees). Eusociality in these families is sometimes managed by a set of
422:
414:
398:
394:
382:
238:
Egg-layers and worker-like individuals among adult females (division of labor)
199:
181:
92:
4992:
4317:
4277:
4238:
3902:
3717:
3709:
3681:
3664:
2211:
1951:
1881:
1497:
species such as Hymenoptera, because of their unusual relatedness structure.
7222:
6862:
6525:
5826:
5821:
5759:
5697:
4900:
4883:
4780:
4576:
3389:
3353:
2109:
1818:
1679:
1623:
1619:
1611:
1150:
1057:
968:
939:
826:
788:
718:
618:
534:
357:
289:
216:
96:
68:
64:
60:
37:
5503:
5448:
5188:
5110:
5051:
4909:
4788:
4702:
4633:
4538:
4335:
4186:
4145:
4104:
4055:
4037:
3998:
3955:
3920:
3811:
3762:
3632:
3590:
3572:
3557:"Ecology, longevity and naked mole-rats: confounding effects of sociality?"
3407:
3277:
3234:
3216:
3139:
2661:
2600:
2539:
2369:
2285:
2266:
2219:
2128:
1969:
409:) contain four tribes of varying degrees of sociality: the highly eusocial
17:
5382:
4584:
4555:; Hare, Hope (1976). "Haplodiploidy and the evolution of social insects".
4481:
3533:
3438:
3090:
2811:
2377:
1388:
877:
7151:
7113:
6845:
6787:
6463:
6114:
6109:
6084:
5983:
5929:
5771:
5764:
5407:
3421:
Burda, H. Honeycutt; Begall, S.; Locker-Grutjen, O.; Scharff, A. (2000).
2620:
1715:
1709:
1666:
1641:
1630:
1615:
1233:
1183:
1021:
988:
953:
892:
834:
792:
538:
402:
390:
277:
5563:
5485:
5092:
5033:
4684:
4520:
2485:
6772:
6149:
5847:
5786:
5776:
4943:
3990:
3793:
2847:
2769:
2591:
2431:
1719:
1477:
1473:
1069:
994:
866:
796:
530:
523:
418:
231:
120:
5494:
5145:"Hemimetabolous genomes reveal molecular basis of termite eusociality"
4834:
4811:
4624:
4384:(2016) . "6. Organisms, Groups, and Memes: Replicators or Vehicles?".
2734:
2673:
2671:
1622:) and physiological effects on the reproductive and endocrine system (
721:
are a class of parasitic flatworm, also known as flukes. One species,
6806:
5744:
4600:"Haplodiploidy and the evolution of eusociality: worker reproduction"
3525:
3069:
Stern, D.; Foster, W. (1996). "The evolution of soldiers in aphids".
3047:
2203:
1610:. Queen mandibular glands produce a mixture of five compounds, three
1047:
978:
945:
777:
622:
591:
586:
406:
378:
293:
144:
132:
100:
88:
72:
56:
4739:
4720:
2942:
2884:
2761:
1986:
Michener, Charles D. (1969). "Comparative Social Behavior of Bees".
655:
Eusociality has evolved in three different lineages in the colonial
450:
71:
from other individuals), overlapping generations within a colony of
4615:
4452:(20 March 1964). "The Genetical Evolution of Social Behaviour II".
2616:"The antiquity and evolutionary history of social behavior in bees"
595:
leaves. Its soldier caste defends the colony in its gall fortress.
7048:
6052:
4425:(20 March 1964). "The Genetical Evolution of Social Behaviour I".
2250:"Evolution of sociality in a primitively eusocial lineage of bees"
1635:
1463:
1373:
1157:
958:
906:
876:
743:
608:
573:
518:
449:
410:
325:
247:
172:
158:
127:
31:
4721:"One Giant Leap: How Insects Achieved Altruism and Colonial Life"
3561:
Proceedings of the Royal Society of London B: Biological Sciences
2934:
1926:"Recent Advances in Behavioral (Epi)Genetics in Eusocial Insects"
165:, too, may make use of a primitively eusocial division of labor.
6794:
6159:
4071:"Evolutionary justifications for human reproductive limitations"
2901:. In: Biocommunication of Animals. Dortrecht, Springer, 161β190.
2087:
1727:
1509:
Against the supposed benefits of haplodiploidy for eusociality,
604:
582:
353:
297:
116:
6719:
6407:
6403:
5956:
5593:
2748:
Conway, John R. (September 1986). "The Biology of Honey Ants".
2404:
Kent, D. S.; Simpson, J. A. (1992). "Eusociality in the beetle
1691:. The pheromones suppress activity of the endocrine gland, the
496:
workers move outside the nest for colony defense and foraging.
266:
Reproductive division of labor (with or without sterile castes)
5739:
5589:
3026:
Crespi B. J. (1992). "Eusociality in Australian gall thrips".
1292:
1279:
1261:
1087:
377:
While only a moderate percentage of species in bees (families
349:
345:
112:
108:
63:. It is defined by the following characteristics: cooperative
2018:
Gadagkar, Raghavendra (1993). "And now... eusocial thrips!".
1682:
from the queen's body alerts those workers of her dominance.
5527:"'Considering Her Ways': In(ter)secting matriarchal utopias"
2899:
Termite Communication During Different Behavioral Activities
2171:
2169:
842:
theory, which grounded Wilson's argument, and because human
6715:
5952:
1868:; Yanega, Douglas (1995). "The Definition of Eusociality".
2040:(1971). "3 The Social Wasps; 4 The Ants; 6 The Termites".
853:. This would mean that humans sometimes exhibit a type of
344:
contains the largest group of eusocial insects, including
4598:
Alpedrinha, JoΓ£o; West, Stuart A.; Gardner, Andy (2013).
3423:"Are naked and common mole-rats eusocial and if so, why?"
2176:
Burns, K. C.; Hutton, Ian; Shepherd, Lara (14 May 2021).
180:
introduced the term "eusocial" after studying nesting in
5579:
2957:"Science: The Australian beetle that behaves like a bee"
2453:
HΓΆlldobler, B. (1990). "8 Caste and Division of Labor".
2073:
2071:
2069:
421:(bumble bees), and the mostly solitary or weakly social
4026:
Proceedings of the Royal Society B: Biological Sciences
3729:
3727:
3372:
Metz, Daniel C. G.; Hechinger, Ryan F. (30 July 2024).
3205:
Proceedings of the Royal Society B: Biological Sciences
2013:
2011:
1981:
1979:
1648:
Among ants, the queen pheromone system of the fire ant
589:(larva on left, adult on right) with galls (centre) on
241:
The overlap of generations (mother and adult offspring)
1505:
Argument that haplodiploidy does not favor eusociality
211:, with individual castes being analogous to different
27:
Highest level of animal sociality a species can attain
5006:
Nowak, M. A.; Tarnita, C. E.; Wilson, E. O. (2010).
2308:
Annual Review of Ecology, Evolution, and Systematics
562:
native to Australia, and is the first beetle (order
244:
Cooperative work on the cells of the bees' honeycomb
131:
exert dominance by fighting. There are two eusocial
7160:
7122:
7077:
7029:
6897:
6753:
6629:
6564:
6441:
6337:
6316:
6195:
5990:
5877:
5856:
5727:
5627:
5580:
International Union for the Study of Social Insects
4888:
Cold Spring Harbor Symposia on Quantitative Biology
1804:
International Union for the Study of Social Insects
1587:the honeybee from reverting to the solitary state.
468:. For example, members of the sterile caste of the
405:. The corbiculate bees (subfamily Apinae of family
300:may exhibit a primitive form of eusocial behavior.
7233:Task allocation and partitioning of social insects
4351:Termites: evolution, sociality, symbioses, ecology
3734:Foster, Kevin R.; Ratnieks, Francis L. W. (2005).
2041:
1837:Task allocation and partitioning of social insects
1759:imagines a human eusocial organisation founded in
4444:
4442:
3604:Foster, Kevin R.; Ratnieks, Francis L.W. (2005).
1489:the fraction of genes that they share, i.e. when
639:(sterile soldier castes being clones produced by
5282:
5280:
5278:
5276:
4965:Shell, Wyatt A.; Rehan, Sandra M. (2017-07-24).
3291:Macdonald, K.S.; Rios, R.; Duffy, J. E. (2006).
3186:Genes, Behaviors and Evolution of Social Insects
2140:
2138:
5274:
5272:
5270:
5268:
5266:
5264:
5262:
5260:
5258:
5256:
3378:Proceedings of the National Academy of Sciences
3342:Proceedings of the National Academy of Sciences
756:Among mammals, two species in the rodent group
4921:
4919:
4495:QuiΓ±ones, AndrΓ©s E.; Pen, Ido (23 June 2017).
3823:
3821:
1672:Similar mechanisms exist in the eusocial wasp
1450:Argument that haplodiploidy favors eusociality
276:Eusociality was then discovered in a group of
198:, who used it to describe nesting behavior in
194:The term "eusocial" was introduced in 1966 by
123:). A colony has caste differences: queens and
6731:
6419:
6324:Association for the Study of Animal Behaviour
5968:
5605:
4752:
4750:
4075:Journal of Assisted Reproduction and Genetics
2614:Cardinal, Sophie; Danforth, Bryan N. (2011).
2297:
2295:
2048:. Cambridge, Massachusetts: Belknap Press of
8:
5216:
5214:
5212:
5210:
5208:
5206:
5204:
5202:
5200:
5198:
4211:; Hodge, S. J.; Flower, T. P. (2008-09-01).
4118:Jetz, Walter; Rubenstein, Dustin R. (2011).
3367:
3365:
3363:
2343:
2341:
2339:
1893:
1891:
1860:
1858:
1856:
1854:
1852:
393:) are eusocial, nearly all species of ants (
5396:
5394:
5392:
4882:Toth, A. L.; Robinson, G. E. (2009-01-01).
2399:
2397:
2395:
825:Scientists have debated whether humans are
6738:
6724:
6716:
6426:
6412:
6404:
6329:International Society for Applied Ethology
5975:
5961:
5953:
5612:
5598:
5590:
5067:"Inclusive fitness theory and eusociality"
4159:Rosenbaum, Stacy; Gettler, Lee T. (2018).
59:) is the highest level of organization of
5493:
5438:
5428:
5178:
5100:
5041:
4982:
4899:
4833:
4738:
4692:
4623:
4528:
4325:
4276:
4176:
4135:
4094:
4045:
3910:
3892:
3875:Kramer, Jos; Meunier, JoΓ«l (2016-04-28).
3801:
3680:
3658:
3656:
3580:
3555:Williams, S. A.; Shattuck, M. R. (2015).
3397:
3316:
3224:
2651:
2641:
2590:
2580:
2563:"Evolutionary History of the Hymenoptera"
2327:
2275:
2265:
2118:
2108:
1959:
1436:quantified a century later with his 1964
887:may display a simple form of eusociality.
36:Co-operative brood rearing, seen here in
4810:Cahan, S. H.; Gardner-Morse, E. (2013).
3698:Personality and Social Psychology Review
2248:Danforth, Bryan N. (December 26, 2001).
2178:"Primitive eusociality in a land plant?"
2081:; HΓΆlldobler, Bert (20 September 2005).
1734:honey bee larvae are initially fed with
3199:Duffy, J. E.; Macdonald, K. S. (2010).
2785:Smithsonian Tropical Research Institute
1848:
417:(stingless bees), primitively eusocial
2304:"Evolution of eusociality in termites"
2147:"These Plants Act Like Bees in a Hive"
2083:"Eusociality: Origin and Consequences"
6841:Patterns of self-organization in ants
3851:"The False Allure of Group Selection"
2897:Costa-Leonardo AM, Haifig I. (2014).
1809:Patterns of self-organization in ants
795:have eusocial tendencies, especially
7:
6686:
5065:Abbot, Patrick; et al. (2011).
6479:Evolutionary models of food sharing
3427:Behavioral Ecology and Sociobiology
2002:10.1146/annurev.en.14.010169.001503
1944:10.1146/annurev-genet-120116-024456
1445:Inclusive fitness and haplodiploidy
1255:
1224:
1189:
1120:
1110:
1063:
1053:
1017:
984:
974:
964:
356:βdivided into castes: reproductive
6851:symmetry breaking of escaping ants
5340:10.1111/j.1365-3113.1987.tb00213.x
4261:"Forum: The eusociality continuum"
3132:10.1111/j.0014-3820.2000.tb00053.x
3083:10.1111/j.1469-185x.1996.tb00741.x
2145:Preston, Elizabeth (2 July 2021).
310:Sexual selection in social insects
87:Eusociality has evolved among the
25:
3743:Trends in Ecology & Evolution
3613:Trends in Ecology & Evolution
2520:Current Opinion in Insect Science
288:Most eusocial societies exist in
6888:
6697:
6685:
6674:
6673:
6388:
6387:
5718:
5239:10.1046/j.1365-3032.1999.00153.x
4655:Nowak, Martin; Tarnita, Corina;
3318:10.1111/j.1366-9516.2005.00213.x
3270:10.1111/j.1461-0248.2012.01788.x
2350:Proceedings of the Royal Society
1907:The Indian Journal of Entomology
1387:
1296:
1268:
1238:
1204:
1160:
1136:
1076:
1033:
997:
40:, is a condition of eusociality.
6613:Sociobiology: The New Synthesis
5922:Sociobiology: The New Synthesis
2320:10.1146/annurev.ecolsys.28.1.27
1814:Reciprocity (social psychology)
1491:Cost < relatedness * Benefit
1419:gene-centered view of evolution
846:is not divided between castes.
103:. It is most widespread in the
75:, and a division of labor into
6033:Bee learning and communication
5585:Eusociality in naked mole-rats
5150:Nature Ecology & Evolution
5008:"The evolution of eusociality"
4661:"The evolution of eusociality"
4454:Journal of Theoretical Biology
4428:Journal of Theoretical Biology
4298:Plant Signaling & Behavior
4292:Burns, Kevin C. (2021-11-02).
3475:. Springer. pp. 207β223.
2408:(Coleoptera: Curculionidae)".
1799:Evolutionarily stable strategy
1767:by E. O. Wilson and others in
1753:'s 2003 science fiction novel
1:
5303:10.1146/annurev.ento.30.1.319
4310:10.1080/15592324.2021.1961063
4231:10.1016/j.anbehav.2008.03.015
4178:10.1016/j.physbeh.2017.12.025
3894:10.12688/f1000research.8018.1
2723:Australian Journal of Zoology
1695:, stopping it from secreting
1640:Physiology of eusociality in
157:and others have claimed that
6812:Mixed-species foraging flock
6763:Agent-based model in biology
5525:Murphy, Graham (July 2008).
5430:10.1371/journal.pone.0010674
5375:10.1016/0003-2697(81)90575-3
4930:(Hymenoptera: Halictidae)".
4474:10.1016/0022-5193(64)90039-6
3736:"A new eusocial vertebrate?"
3606:"A new eusocial vertebrate?"
3481:10.1007/978-3-540-75957-7_10
2750:The American Biology Teacher
2643:10.1371/journal.pone.0021086
2457:. Cambridge, Massachusetts:
776:), both of which are highly
7059:Particle swarm optimization
6704:Evolutionary biology portal
5688:Sexual selection in insects
5290:Annual Review of Entomology
3473:Ecology of Social Evolution
3297:Diversity and Distributions
2872:Canadian Journal of Zoology
1989:Annual Review of Entomology
292:, while a few are found in
7280:
6768:Collective animal behavior
5840:Austroplatypus incompertus
5668:Identity in social insects
4355:Kluwer Academic Publishers
3755:10.1016/j.tree.2005.05.005
3625:10.1016/j.tree.2005.05.005
3167:Bulletin of Marine Science
2532:10.1016/j.cois.2019.04.006
2406:Austroplatypus incompertus
1591:Physiology and development
1554:
1453:
1201:Austroplatypus incompertus
937:
926:
818:
555:Austroplatypus incompertus
314:Identity in social insects
307:
6886:
6669:
6383:
6090:Evolutionary neuroscience
5716:
5171:10.1038/s41559-017-0459-1
4984:10.1007/s13592-017-0527-1
4869:10.1017/s0952836902000481
4165:Physiology & Behavior
4137:10.1016/j.cub.2010.11.075
4087:10.1007/s10815-018-1285-3
3979:Evolutionary Anthropology
3948:10.1007/s12110-013-9182-z
3887:: F1000 Faculty Revβ776.
3005:10.1007/s10144-005-0218-z
2914:Journal of Animal Ecology
2700:10.1007/s00040-010-0080-7
2582:10.1016/j.cub.2017.01.027
2474:Annales Zoologici Fennici
1931:Annual Review of Genetics
1563:between-group competition
1377:
1288:
1266:
1253:
1229:
1222:
1197:
1187:
1154:
1128:
1118:
1108:
1074:
1061:
1051:
1025:
1015:
992:
982:
972:
934:Phylogenetic distribution
915:specialized for tasks, a
466:self-sacrificing behavior
272:Cooperative care of young
7097:Self-propelled particles
6043:Behavioral endocrinology
5635:Evolution of eusociality
5227:Physiological Entomology
4412:v7, pp. 1β16, and 17-52.
4369:On the Origin of Species
3710:10.1177/1088868311430834
3682:10.1525/bio.2012.62.11.8
3663:Gintis, Herbert (2012).
2050:Harvard University Press
1424:On the Origin of Species
929:Evolution of eusociality
570:In gall-inducing insects
502:Lasioglossum aeneiventre
67:care (including care of
7178:Collective intelligence
7044:Ant colony optimization
6499:Male warrior hypothesis
6469:Evolutionary psychology
6459:Dual inheritance theory
6238:IrenΓ€us Eibl-Eibesfeldt
6018:Animal sexual behaviour
5898:Charles Duncan Michener
5531:Science Fiction Studies
5363:Analytical Biochemistry
4901:10.1101/sqb.2009.74.026
4781:10.1126/science.1156108
4604:The American Naturalist
4577:10.1126/science.1108197
4390:Oxford University Press
4069:Lubinsky, Mark (2018).
3390:10.1073/pnas.2400953121
3354:10.1073/pnas.2414228121
2110:10.1073/pnas.0505858102
1830:Bee colony optimization
1824:Ant colony optimization
1631:emergence from the pupa
508:Lasioglossum malachurum
269:Overlapping generations
7198:Microbial intelligence
6858:Shoaling and schooling
6177:Tool use by non-humans
6130:Philosophical ethology
6075:Comparative psychology
6023:Animal welfare science
4386:The Extended Phenotype
4278:10.1093/beheco/6.1.102
4038:10.1098/rspb.2014.2808
3782:Psychological Bulletin
3573:10.1098/rspb.2014.1664
3217:10.1098/rspb.2009.1483
2370:10.1098/rspb.1994.0071
2302:Thorne, B. L. (1997).
2267:10.1073/pnas.012387999
1882:10.1093/beheco/6.1.109
1645:
1578:Reversal to solitarity
1481:
1384:Platycerium bifurcatum
898:Platycerium bifurcatum
888:
884:Platycerium bifurcatum
863:some non-eusocial bees
753:
596:
527:
460:
444:Tetragonula carbonaria
337:
256:
221:multicellular organism
191:
41:
6474:Evolution of morality
5903:The Bees of the World
5319:Systematic Entomology
4814:Pogonomyrmex barbatus
4501:Nature Communications
3439:10.1007/s002650050669
3161:Duffy, J. E. (1998).
2812:10.1007/s000400050063
1639:
1555:Further information:
1467:
1454:Further information:
938:Further information:
880:
819:Further information:
766:Heterocephalus glaber
747:
614:Pemphigus spyrothecae
601:gall-inducing insects
577:
522:
488:Scaptotrigona postica
453:
333:Iridomyrmex purpureus
329:
251:
187:Halictus latisignatus
176:
35:
7218:Spatial organization
7183:Decentralised system
7021:Sea turtle migration
6875:Swarming (honey bee)
6454:Challenge hypothesis
6449:Behavioural genetics
6283:William Homan Thorpe
6048:Behavioural genetics
6008:Animal consciousness
6003:Animal communication
4851:Cryptomys damarensis
4392:. pp. 147β178.
4271:(1): 102β108. 1995.
4209:Clutton-Brock, T. H.
2044:The Insect Societies
1901:(1 September 1966).
1899:Batra, Suzanne W. T.
1662:Oecophylla longinoda
1551:Multilevel selection
1529:Damaraland mole-rats
1522:Evolutionary ecology
787:Some mammals in the
637:asexual reproduction
119:) and in Blattodea (
7193:Group size measures
6755:Biological swarming
6504:Reciprocal altruism
6038:Behavioural ecology
5938:Journey to the Ants
5486:10.1038/nature10093
5478:2011Natur.473..478K
5421:2010PLoSO...510674H
5332:1987SysEn..12..413C
5163:2018NatEE...2..557H
5093:10.1038/nature09831
5085:2011Natur.471E...1A
5034:10.1038/nature09205
5026:2010Natur.466.1057N
5020:(7310): 1057β1062.
4928:Halictus rubicundus
4773:2008Sci...320.1213H
4767:(5880): 1213β1216.
4685:10.1038/nature09205
4677:2010Natur.466.1057N
4671:(7310): 1057β1062.
4569:1976Sci...191..249T
4521:10.1038/ncomms15920
4513:2017NatCo...815920Q
4466:1964JThBi...7...17H
3518:1996Natur.380..619O
3309:2006DivDi..12..165M
3262:2012EcolL..15..704H
3040:1992Natur.359..724C
2997:2005PopEc..47..127A
2927:1987JAnEc..56.1069A
2634:2011PLoSO...621086C
2461:. pp. 298β318.
2424:1992NW.....79...86K
2411:Naturwissenschaften
2362:1994RSPSB.256..203S
2196:2021Ecol..102E3373B
2101:2005PNAS..10213367W
2095:(38): 13367β13371.
859:helpers at the nest
857:behavior known as "
770:Damaraland mole-rat
740:In nonhuman mammals
371:Polistes versicolor
228:Charles D. Michener
141:Damaraland mole-rat
7254:Behavioral ecology
7208:Predator satiation
7069:Swarm (simulation)
7064:Swarm intelligence
7039:Agent-based models
6870:Swarming behaviour
6592:Sarah Blaffer Hrdy
6521:Sex and psychology
6367:Behavioral Ecology
6288:Nikolaas Tinbergen
6080:Emotion in animals
6058:Cognitive ethology
5223:Solenopsis invicta
4944:10.1007/BF01240705
4856:Journal of Zoology
4821:Journal of Zoology
4659:(26 August 2010).
4553:Trivers, Robert L.
4265:Behavioral Ecology
4032:(1803): 20142808.
3991:10.1002/evan.21382
3794:10.1037/bul0000123
3567:(1802): 20141664.
3071:Biological Reviews
2984:Population Ecology
2848:10.1007/BF01338830
2680:Apis andreniformis
2432:10.1007/BF01131810
1870:Behavioral Ecology
1866:Crespi, Bernard J.
1765:Groping for Groups
1746:In popular culture
1651:Solenopsis invicta
1646:
1482:
948:, as shown in the
889:
881:The staghorn fern
844:reproductive labor
801:Suricata suricatta
774:Fukomys damarensis
760:are eusocial, the
754:
724:Haplorchis pumilio
597:
579:Kladothrips rugosa
528:
461:
432:Apis andreniformis
338:
257:
192:
150:Synalpheus regalis
42:
7241:
7240:
7228:Military swarming
7173:Animal navigation
7092:Collective motion
7079:Collective motion
6946:reverse migration
6880:Swarming motility
6713:
6712:
6637:Stephen Jay Gould
6401:
6400:
6293:Jakob von UexkΓΌll
6063:Comfort behaviour
5950:
5949:
5864:Bees in mythology
5551:The Hudson Review
5472:(7348): 478β483.
4835:10.1111/jzo.12071
4717:Wilson, Edward O.
4657:Wilson, Edward O.
4563:(4224): 249β263.
4371:, 1859. Chapter 8
4367:Darwin, Charles.
4357:. pp. 77β93.
4081:(12): 2133β2139.
3788:(12): 1313β1345.
3512:(6575): 619β621.
3490:978-3-540-75956-0
3211:(1681): 575β584.
3034:(6397): 724β726.
2735:10.1071/zo9960143
2356:(1346): 203β209.
2079:Wilson, Edward O.
2038:Wilson, Edward O.
1770:The Hudson Review
1688:Bombus terrestris
1486:inclusive fitness
1456:Inclusive fitness
1438:inclusive fitness
1409:
1408:
1400:
1399:
1363:
1362:
1354:
1353:
1345:
1344:
1336:
1335:
1327:
1326:
1318:
1317:
1309:
1308:
1173:
1172:
1094:
1093:
950:phylogenetic tree
917:division of labor
731:. Interestingly,
413:(honey bees) and
16:(Redirected from
7271:
7054:Crowd simulation
7031:Swarm algorithms
7002:Insect migration
6907:Animal migration
6899:Animal migration
6892:
6817:Mobbing behavior
6740:
6733:
6726:
6717:
6701:
6689:
6688:
6677:
6676:
6658:Not in Our Genes
6647:Richard Lewontin
6509:Sexual selection
6428:
6421:
6414:
6405:
6391:
6390:
6353:Animal Cognition
6346:Animal Behaviour
6298:Wolfgang Wickler
5998:Animal cognition
5977:
5970:
5963:
5954:
5890:The Dancing Bees
5722:
5614:
5607:
5600:
5591:
5568:
5567:
5545:
5539:
5538:
5522:
5516:
5515:
5497:
5459:
5453:
5452:
5442:
5432:
5398:
5387:
5386:
5358:
5352:
5351:
5313:
5307:
5306:
5284:
5251:
5250:
5218:
5193:
5192:
5182:
5140:
5134:
5133:
5121:
5115:
5114:
5104:
5062:
5056:
5055:
5045:
5003:
4997:
4996:
4986:
4962:
4956:
4955:
4932:Insectes Sociaux
4923:
4914:
4913:
4903:
4879:
4873:
4872:
4846:
4840:
4839:
4837:
4807:
4801:
4800:
4754:
4745:
4744:
4742:
4713:
4707:
4706:
4696:
4652:
4646:
4645:
4627:
4595:
4589:
4588:
4549:
4543:
4542:
4532:
4492:
4486:
4485:
4446:
4437:
4436:
4419:
4413:
4410:
4404:
4403:
4382:Dawkins, Richard
4378:
4372:
4365:
4359:
4358:
4346:
4340:
4339:
4329:
4289:
4283:
4282:
4280:
4257:
4251:
4250:
4218:Animal Behaviour
4205:
4199:
4198:
4180:
4156:
4150:
4149:
4139:
4115:
4109:
4108:
4098:
4066:
4060:
4059:
4049:
4017:
4011:
4010:
3974:
3968:
3967:
3931:
3925:
3924:
3914:
3896:
3872:
3866:
3865:
3863:
3861:
3843:
3837:
3836:
3829:Dawkins, Richard
3825:
3816:
3815:
3805:
3773:
3767:
3766:
3740:
3731:
3722:
3721:
3693:
3687:
3686:
3684:
3660:
3651:
3650:
3648:
3647:
3641:
3635:. Archived from
3610:
3601:
3595:
3594:
3584:
3552:
3546:
3545:
3526:10.1038/380619a0
3501:
3495:
3494:
3468:
3462:
3461:
3459:
3458:
3449:. Archived from
3418:
3412:
3411:
3401:
3369:
3358:
3357:
3337:
3331:
3330:
3320:
3288:
3282:
3281:
3245:
3239:
3238:
3228:
3196:
3190:
3189:
3181:
3175:
3174:
3158:
3152:
3151:
3109:
3103:
3102:
3066:
3060:
3059:
3048:10.1038/359724a0
3023:
3017:
3016:
2978:
2972:
2971:
2969:
2968:
2953:
2947:
2946:
2921:(3): 1069β1081.
2908:
2902:
2895:
2889:
2888:
2879:(7): 1259β1266.
2866:
2860:
2859:
2836:Insectes Sociaux
2830:
2824:
2823:
2800:Insectes Sociaux
2795:
2789:
2788:
2780:
2774:
2773:
2745:
2739:
2738:
2718:
2712:
2711:
2688:Insectes Sociaux
2675:
2666:
2665:
2655:
2645:
2611:
2605:
2604:
2594:
2584:
2575:(7): 1013β1018.
2558:
2552:
2551:
2515:
2509:
2508:
2496:
2490:
2489:
2480:(5/6): 531β549.
2469:
2463:
2462:
2450:
2444:
2443:
2401:
2390:
2389:
2345:
2334:
2333:
2331:
2299:
2290:
2289:
2279:
2269:
2245:
2239:
2238:
2236:
2234:
2204:10.1002/ecy.3373
2173:
2164:
2163:
2161:
2159:
2142:
2133:
2132:
2122:
2112:
2075:
2064:
2063:
2047:
2034:
2028:
2027:
2015:
2006:
2005:
1983:
1974:
1973:
1963:
1921:
1915:
1914:
1895:
1886:
1885:
1862:
1794:Dense heterarchy
1703:Other mechanisms
1697:juvenile hormone
1675:Vespula vulgaris
1391:
1300:
1272:
1256:
1242:
1225:
1208:
1190:
1164:
1140:
1121:
1111:
1080:
1064:
1054:
1037:
1018:
1001:
985:
975:
965:
831:Edward O. Wilson
809:Helogale parvula
782:avoid inbreeding
674:S. microneptunus
603:, including the
558:is a species of
482:Synoeca surinama
153:, are eusocial.
21:
7279:
7278:
7274:
7273:
7272:
7270:
7269:
7268:
7244:
7243:
7242:
7237:
7156:
7118:
7073:
7025:
6893:
6884:
6749:
6744:
6714:
6709:
6665:
6625:
6582:Richard Dawkins
6560:
6551:Dunbar's number
6489:Kin recognition
6484:Group selection
6437:
6432:
6402:
6397:
6379:
6333:
6312:
6308:Solly Zuckerman
6248:Karl von Frisch
6233:Richard Dawkins
6218:John B. Calhoun
6203:Patrick Bateson
6191:
6125:Pain in animals
5986:
5981:
5951:
5946:
5911:Bert HΓΆlldobler
5885:Karl von Frisch
5878:Pioneers, works
5873:
5852:
5723:
5714:
5708:Worker policing
5693:Social conflict
5673:Kin recognition
5658:Group selection
5644:Social insects
5623:
5618:
5576:
5571:
5547:
5546:
5542:
5524:
5523:
5519:
5461:
5460:
5456:
5400:
5399:
5390:
5360:
5359:
5355:
5315:
5314:
5310:
5286:
5285:
5254:
5220:
5219:
5196:
5142:
5141:
5137:
5123:
5122:
5118:
5079:(7339): E1βE4.
5064:
5063:
5059:
5005:
5004:
5000:
4964:
4963:
4959:
4925:
4924:
4917:
4881:
4880:
4876:
4848:
4847:
4843:
4809:
4808:
4804:
4756:
4755:
4748:
4740:10.1641/b580106
4715:
4714:
4710:
4654:
4653:
4649:
4597:
4596:
4592:
4551:
4550:
4546:
4494:
4493:
4489:
4450:Hamilton, W. D.
4448:
4447:
4440:
4423:Hamilton, W. D.
4421:
4420:
4416:
4411:
4407:
4400:
4380:
4379:
4375:
4366:
4362:
4348:
4347:
4343:
4291:
4290:
4286:
4259:
4258:
4254:
4207:
4206:
4202:
4158:
4157:
4153:
4124:Current Biology
4117:
4116:
4112:
4068:
4067:
4063:
4019:
4018:
4014:
3976:
3975:
3971:
3933:
3932:
3928:
3874:
3873:
3869:
3859:
3857:
3845:
3844:
3840:
3827:
3826:
3819:
3775:
3774:
3770:
3738:
3733:
3732:
3725:
3695:
3694:
3690:
3675:(11): 987β991.
3662:
3661:
3654:
3645:
3643:
3639:
3608:
3603:
3602:
3598:
3554:
3553:
3549:
3503:
3502:
3498:
3491:
3470:
3469:
3465:
3456:
3454:
3420:
3419:
3415:
3371:
3370:
3361:
3339:
3338:
3334:
3290:
3289:
3285:
3250:Ecology Letters
3247:
3246:
3242:
3198:
3197:
3193:
3183:
3182:
3178:
3160:
3159:
3155:
3111:
3110:
3106:
3068:
3067:
3063:
3025:
3024:
3020:
2980:
2979:
2975:
2966:
2964:
2955:
2954:
2950:
2910:
2909:
2905:
2896:
2892:
2885:10.1139/z00-064
2868:
2867:
2863:
2832:
2831:
2827:
2797:
2796:
2792:
2782:
2781:
2777:
2762:10.2307/4448321
2747:
2746:
2742:
2720:
2719:
2715:
2677:
2676:
2669:
2613:
2612:
2608:
2568:Current Biology
2560:
2559:
2555:
2517:
2516:
2512:
2498:
2497:
2493:
2471:
2470:
2466:
2452:
2451:
2447:
2403:
2402:
2393:
2347:
2346:
2337:
2301:
2300:
2293:
2247:
2246:
2242:
2232:
2230:
2175:
2174:
2167:
2157:
2155:
2144:
2143:
2136:
2077:
2076:
2067:
2060:
2036:
2035:
2031:
2020:Current Science
2017:
2016:
2009:
1985:
1984:
1977:
1923:
1922:
1918:
1897:
1896:
1889:
1864:
1863:
1850:
1846:
1841:
1789:
1780:Brave New World
1748:
1705:
1598:
1593:
1580:
1559:
1557:Group selection
1553:
1524:
1507:
1468:Honey bees are
1462:
1452:
1447:
1415:
1410:
1401:
1364:
1355:
1346:
1337:
1328:
1319:
1310:
1174:
1095:
942:
936:
931:
925:
895:staghorn fern,
891:One plant, the
875:
840:group selection
823:
821:Group selection
817:
805:dwarf mongooses
742:
715:
699:S. cayoneptunus
653:
645:kleptoparasites
641:parthenogenesis
572:
560:ambrosia beetle
551:
517:
364:, more or less
324:
316:
306:
286:
184:bees including
171:
28:
23:
22:
15:
12:
11:
5:
7277:
7275:
7267:
7266:
7261:
7259:Superorganisms
7256:
7246:
7245:
7239:
7238:
7236:
7235:
7230:
7225:
7220:
7215:
7213:Quorum sensing
7210:
7205:
7200:
7195:
7190:
7185:
7180:
7175:
7170:
7164:
7162:
7161:Related topics
7158:
7157:
7155:
7154:
7149:
7147:Swarm robotics
7144:
7139:
7134:
7128:
7126:
7124:Swarm robotics
7120:
7119:
7117:
7116:
7111:
7106:
7105:
7104:
7094:
7089:
7083:
7081:
7075:
7074:
7072:
7071:
7066:
7061:
7056:
7051:
7046:
7041:
7035:
7033:
7027:
7026:
7024:
7023:
7018:
7017:
7016:
7015:
7014:
6999:
6998:
6997:
6992:
6982:
6981:
6980:
6975:
6970:
6965:
6958:Fish migration
6955:
6953:Cell migration
6950:
6949:
6948:
6943:
6936:Bird migration
6933:
6932:
6931:
6929:coded wire tag
6926:
6925:
6924:
6914:
6903:
6901:
6895:
6894:
6887:
6885:
6883:
6882:
6877:
6872:
6867:
6866:
6865:
6855:
6854:
6853:
6848:
6838:
6837:
6836:
6826:
6825:
6824:
6822:feeding frenzy
6814:
6809:
6804:
6803:
6802:
6792:
6791:
6790:
6785:
6775:
6770:
6765:
6759:
6757:
6751:
6750:
6745:
6743:
6742:
6735:
6728:
6720:
6711:
6710:
6708:
6707:
6695:
6683:
6670:
6667:
6666:
6664:
6663:
6662:
6661:
6649:
6644:
6639:
6633:
6631:
6627:
6626:
6624:
6623:
6618:
6617:
6616:
6604:
6599:
6594:
6589:
6587:Daniel Dennett
6584:
6579:
6574:
6568:
6566:
6562:
6561:
6559:
6558:
6553:
6548:
6547:
6546:
6540:
6539:
6538:
6523:
6518:
6517:
6516:
6506:
6501:
6496:
6491:
6486:
6481:
6476:
6471:
6466:
6461:
6456:
6451:
6445:
6443:
6439:
6438:
6433:
6431:
6430:
6423:
6416:
6408:
6399:
6398:
6396:
6395:
6384:
6381:
6380:
6378:
6377:
6370:
6363:
6360:Animal Welfare
6356:
6349:
6341:
6339:
6335:
6334:
6332:
6331:
6326:
6320:
6318:
6314:
6313:
6311:
6310:
6305:
6300:
6295:
6290:
6285:
6280:
6275:
6273:Desmond Morris
6270:
6265:
6260:
6255:
6250:
6245:
6240:
6235:
6230:
6228:Marian Dawkins
6225:
6223:Charles Darwin
6220:
6215:
6210:
6205:
6199:
6197:
6193:
6192:
6190:
6189:
6184:
6179:
6174:
6169:
6168:
6167:
6162:
6157:
6152:
6142:
6137:
6132:
6127:
6122:
6117:
6112:
6107:
6105:Human ethology
6102:
6097:
6092:
6087:
6082:
6077:
6072:
6071:
6070:
6060:
6055:
6050:
6045:
6040:
6035:
6030:
6025:
6020:
6015:
6013:Animal culture
6010:
6005:
6000:
5994:
5992:
5988:
5987:
5982:
5980:
5979:
5972:
5965:
5957:
5948:
5947:
5945:
5944:
5943:
5942:
5934:
5926:
5913:
5908:
5907:
5906:
5895:
5894:
5893:
5881:
5879:
5875:
5874:
5872:
5871:
5866:
5860:
5858:
5854:
5853:
5851:
5850:
5845:
5844:
5843:
5831:
5830:
5829:
5819:
5818:
5817:
5805:
5804:
5803:
5791:
5790:
5789:
5784:
5782:Dwarf mongoose
5779:
5769:
5768:
5767:
5762:
5757:
5752:
5747:
5742:
5731:
5729:
5725:
5724:
5717:
5715:
5713:
5712:
5711:
5710:
5705:
5700:
5695:
5690:
5685:
5680:
5675:
5670:
5665:
5660:
5655:
5650:
5642:
5637:
5631:
5629:
5625:
5624:
5619:
5617:
5616:
5609:
5602:
5594:
5588:
5587:
5582:
5575:
5574:External links
5572:
5570:
5569:
5558:(4): 652β658.
5540:
5517:
5454:
5388:
5369:(2): 372β375.
5353:
5326:(4): 413β431.
5308:
5252:
5233:(4): 370β376.
5194:
5157:(3): 557β566.
5135:
5116:
5057:
4998:
4957:
4938:(2): 169β180.
4915:
4874:
4863:(4): 445β448.
4841:
4802:
4746:
4719:(2008-01-01).
4708:
4647:
4616:10.1086/671994
4610:(4): 421β438.
4590:
4544:
4487:
4438:
4414:
4405:
4399:978-0198788911
4398:
4373:
4360:
4341:
4284:
4252:
4225:(3): 689β700.
4200:
4171:(Pt A): 1β11.
4151:
4110:
4061:
4012:
3985:(6): 294β302.
3969:
3942:(4): 375β401.
3926:
3867:
3847:Pinker, Steven
3838:
3817:
3768:
3749:(7): 363β364.
3723:
3704:(3): 233β261.
3688:
3652:
3619:(7): 363β364.
3596:
3547:
3496:
3489:
3463:
3433:(5): 293β303.
3413:
3359:
3332:
3303:(2): 165β178.
3283:
3256:(7): 704β713.
3240:
3191:
3176:
3153:
3126:(2): 503β516.
3104:
3061:
3018:
2991:(2): 127β136.
2973:
2948:
2903:
2890:
2861:
2825:
2806:(4): 435β447.
2790:
2775:
2756:(6): 335β343.
2740:
2713:
2694:(3): 251β255.
2667:
2606:
2553:
2510:
2491:
2464:
2445:
2391:
2335:
2291:
2260:(1): 286β290.
2240:
2165:
2152:New York Times
2134:
2065:
2058:
2029:
2007:
1975:
1938:(1): 489β510.
1916:
1887:
1847:
1845:
1842:
1840:
1839:
1834:
1833:
1832:
1827:
1816:
1811:
1806:
1801:
1796:
1790:
1788:
1785:
1751:Stephen Baxter
1747:
1744:
1704:
1701:
1693:corpus allatum
1607:Apis mellifera
1597:
1594:
1592:
1589:
1579:
1576:
1552:
1549:
1523:
1520:
1511:Robert Trivers
1506:
1503:
1451:
1448:
1446:
1443:
1414:
1411:
1407:
1406:
1403:
1402:
1398:
1397:
1394:
1393:
1376:
1370:
1369:
1366:
1365:
1361:
1360:
1357:
1356:
1352:
1351:
1348:
1347:
1343:
1342:
1339:
1338:
1334:
1333:
1330:
1329:
1325:
1324:
1321:
1320:
1316:
1315:
1312:
1311:
1307:
1306:
1303:
1302:
1287:
1284:
1283:
1275:
1274:
1265:
1254:
1252:
1249:
1248:
1245:
1244:
1228:
1223:
1221:
1215:
1214:
1211:
1210:
1196:
1188:
1186:
1180:
1179:
1176:
1175:
1171:
1170:
1167:
1166:
1153:
1147:
1146:
1143:
1142:
1127:
1119:
1117:
1109:
1107:
1101:
1100:
1097:
1096:
1092:
1091:
1083:
1082:
1073:
1062:
1060:
1052:
1050:
1044:
1043:
1040:
1039:
1024:
1016:
1014:
1008:
1007:
1004:
1003:
991:
983:
981:
973:
971:
963:
946:animal kingdom
935:
932:
927:Main article:
924:
921:
874:
871:
816:
813:
762:naked mole-rat
750:naked mole-rat
741:
738:
714:
713:In trematodes
711:
683:S. elizabethae
679:S. filidigitus
652:
651:In crustaceans
649:
571:
568:
550:
547:
516:
513:
323:
322:In hymenoptera
320:
305:
302:
285:
282:
274:
273:
270:
267:
246:
245:
242:
239:
209:superorganisms
170:
167:
137:naked mole-rat
82:superorganisms
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
7276:
7265:
7262:
7260:
7257:
7255:
7252:
7251:
7249:
7234:
7231:
7229:
7226:
7224:
7221:
7219:
7216:
7214:
7211:
7209:
7206:
7204:
7201:
7199:
7196:
7194:
7191:
7189:
7186:
7184:
7181:
7179:
7176:
7174:
7171:
7169:
7166:
7165:
7163:
7159:
7153:
7150:
7148:
7145:
7143:
7140:
7138:
7135:
7133:
7130:
7129:
7127:
7125:
7121:
7115:
7112:
7110:
7107:
7103:
7100:
7099:
7098:
7095:
7093:
7090:
7088:
7087:Active matter
7085:
7084:
7082:
7080:
7076:
7070:
7067:
7065:
7062:
7060:
7057:
7055:
7052:
7050:
7047:
7045:
7042:
7040:
7037:
7036:
7034:
7032:
7028:
7022:
7019:
7013:
7010:
7009:
7008:
7005:
7004:
7003:
7000:
6996:
6993:
6991:
6988:
6987:
6986:
6983:
6979:
6976:
6974:
6971:
6969:
6966:
6964:
6963:diel vertical
6961:
6960:
6959:
6956:
6954:
6951:
6947:
6944:
6942:
6939:
6938:
6937:
6934:
6930:
6927:
6923:
6920:
6919:
6918:
6915:
6913:
6910:
6909:
6908:
6905:
6904:
6902:
6900:
6896:
6891:
6881:
6878:
6876:
6873:
6871:
6868:
6864:
6861:
6860:
6859:
6856:
6852:
6849:
6847:
6844:
6843:
6842:
6839:
6835:
6832:
6831:
6830:
6827:
6823:
6820:
6819:
6818:
6815:
6813:
6810:
6808:
6805:
6801:
6800:herd behavior
6798:
6797:
6796:
6793:
6789:
6786:
6784:
6781:
6780:
6779:
6776:
6774:
6771:
6769:
6766:
6764:
6761:
6760:
6758:
6756:
6752:
6748:
6741:
6736:
6734:
6729:
6727:
6722:
6721:
6718:
6706:
6705:
6700:
6696:
6694:
6693:
6684:
6682:
6681:
6672:
6671:
6668:
6660:
6659:
6655:
6654:
6653:
6650:
6648:
6645:
6643:
6640:
6638:
6635:
6634:
6632:
6628:
6622:
6621:Robert Wright
6619:
6615:
6614:
6610:
6609:
6608:
6605:
6603:
6602:Frans de Waal
6600:
6598:
6597:Steven Pinker
6595:
6593:
6590:
6588:
6585:
6583:
6580:
6578:
6575:
6573:
6572:Anne Campbell
6570:
6569:
6567:
6563:
6557:
6554:
6552:
6549:
6544:
6541:
6537:
6534:
6533:
6532:
6529:
6528:
6527:
6524:
6522:
6519:
6515:
6512:
6511:
6510:
6507:
6505:
6502:
6500:
6497:
6495:
6494:Kin selection
6492:
6490:
6487:
6485:
6482:
6480:
6477:
6475:
6472:
6470:
6467:
6465:
6462:
6460:
6457:
6455:
6452:
6450:
6447:
6446:
6444:
6440:
6436:
6429:
6424:
6422:
6417:
6415:
6410:
6409:
6406:
6394:
6386:
6385:
6382:
6376:
6375:
6371:
6369:
6368:
6364:
6362:
6361:
6357:
6355:
6354:
6350:
6348:
6347:
6343:
6342:
6340:
6336:
6330:
6327:
6325:
6322:
6321:
6319:
6315:
6309:
6306:
6304:
6301:
6299:
6296:
6294:
6291:
6289:
6286:
6284:
6281:
6279:
6278:Thomas Sebeok
6276:
6274:
6271:
6269:
6268:Konrad Lorenz
6266:
6264:
6263:Julian Huxley
6261:
6259:
6258:Heini Hediger
6256:
6254:
6251:
6249:
6246:
6244:
6241:
6239:
6236:
6234:
6231:
6229:
6226:
6224:
6221:
6219:
6216:
6214:
6211:
6209:
6206:
6204:
6201:
6200:
6198:
6194:
6188:
6187:Zoomusicology
6185:
6183:
6180:
6178:
6175:
6173:
6170:
6166:
6163:
6161:
6158:
6156:
6153:
6151:
6148:
6147:
6146:
6143:
6141:
6138:
6136:
6133:
6131:
6128:
6126:
6123:
6121:
6120:Neuroethology
6118:
6116:
6113:
6111:
6108:
6106:
6103:
6101:
6098:
6096:
6093:
6091:
6088:
6086:
6083:
6081:
6078:
6076:
6073:
6069:
6066:
6065:
6064:
6061:
6059:
6056:
6054:
6051:
6049:
6046:
6044:
6041:
6039:
6036:
6034:
6031:
6029:
6028:Anthrozoology
6026:
6024:
6021:
6019:
6016:
6014:
6011:
6009:
6006:
6004:
6001:
5999:
5996:
5995:
5993:
5989:
5985:
5978:
5973:
5971:
5966:
5964:
5959:
5958:
5955:
5940:
5939:
5935:
5932:
5931:
5927:
5924:
5923:
5919:
5918:
5917:
5914:
5912:
5909:
5904:
5901:
5900:
5899:
5896:
5891:
5888:
5887:
5886:
5883:
5882:
5880:
5876:
5870:
5867:
5865:
5862:
5861:
5859:
5855:
5849:
5846:
5842:
5841:
5837:
5836:
5835:
5832:
5828:
5825:
5824:
5823:
5820:
5816:
5815:
5811:
5810:
5809:
5806:
5802:
5801:
5797:
5796:
5795:
5792:
5788:
5785:
5783:
5780:
5778:
5775:
5774:
5773:
5770:
5766:
5763:
5761:
5758:
5756:
5753:
5751:
5748:
5746:
5743:
5741:
5738:
5737:
5736:
5733:
5732:
5730:
5726:
5721:
5709:
5706:
5704:
5701:
5699:
5696:
5694:
5691:
5689:
5686:
5684:
5681:
5679:
5678:Kin selection
5676:
5674:
5671:
5669:
5666:
5664:
5663:Haplodiploidy
5661:
5659:
5656:
5654:
5651:
5649:
5646:
5645:
5643:
5641:
5638:
5636:
5633:
5632:
5630:
5626:
5622:
5615:
5610:
5608:
5603:
5601:
5596:
5595:
5592:
5586:
5583:
5581:
5578:
5577:
5573:
5565:
5561:
5557:
5553:
5552:
5544:
5541:
5537:(2): 266β280.
5536:
5532:
5528:
5521:
5518:
5513:
5509:
5505:
5501:
5496:
5491:
5487:
5483:
5479:
5475:
5471:
5467:
5466:
5458:
5455:
5450:
5446:
5441:
5436:
5431:
5426:
5422:
5418:
5415:(5): e10674.
5414:
5410:
5409:
5404:
5397:
5395:
5393:
5389:
5384:
5380:
5376:
5372:
5368:
5364:
5357:
5354:
5349:
5345:
5341:
5337:
5333:
5329:
5325:
5321:
5320:
5312:
5309:
5304:
5300:
5296:
5292:
5291:
5283:
5281:
5279:
5277:
5275:
5273:
5271:
5269:
5267:
5265:
5263:
5261:
5259:
5257:
5253:
5248:
5244:
5240:
5236:
5232:
5228:
5224:
5217:
5215:
5213:
5211:
5209:
5207:
5205:
5203:
5201:
5199:
5195:
5190:
5186:
5181:
5176:
5172:
5168:
5164:
5160:
5156:
5152:
5151:
5146:
5139:
5136:
5131:
5127:
5120:
5117:
5112:
5108:
5103:
5098:
5094:
5090:
5086:
5082:
5078:
5074:
5073:
5068:
5061:
5058:
5053:
5049:
5044:
5039:
5035:
5031:
5027:
5023:
5019:
5015:
5014:
5009:
5002:
4999:
4994:
4990:
4985:
4980:
4976:
4972:
4968:
4961:
4958:
4953:
4949:
4945:
4941:
4937:
4933:
4929:
4922:
4920:
4916:
4911:
4907:
4902:
4897:
4893:
4889:
4885:
4878:
4875:
4870:
4866:
4862:
4858:
4857:
4852:
4845:
4842:
4836:
4831:
4827:
4823:
4822:
4817:
4815:
4806:
4803:
4798:
4794:
4790:
4786:
4782:
4778:
4774:
4770:
4766:
4762:
4761:
4753:
4751:
4747:
4741:
4736:
4732:
4728:
4727:
4722:
4718:
4712:
4709:
4704:
4700:
4695:
4690:
4686:
4682:
4678:
4674:
4670:
4666:
4662:
4658:
4651:
4648:
4643:
4639:
4635:
4631:
4626:
4621:
4617:
4613:
4609:
4605:
4601:
4594:
4591:
4586:
4582:
4578:
4574:
4570:
4566:
4562:
4558:
4554:
4548:
4545:
4540:
4536:
4531:
4526:
4522:
4518:
4514:
4510:
4506:
4502:
4498:
4491:
4488:
4483:
4479:
4475:
4471:
4467:
4463:
4459:
4455:
4451:
4445:
4443:
4439:
4434:
4430:
4429:
4424:
4418:
4415:
4409:
4406:
4401:
4395:
4391:
4387:
4383:
4377:
4374:
4370:
4364:
4361:
4356:
4352:
4345:
4342:
4337:
4333:
4328:
4323:
4319:
4315:
4311:
4307:
4303:
4299:
4295:
4288:
4285:
4279:
4274:
4270:
4266:
4262:
4256:
4253:
4248:
4244:
4240:
4236:
4232:
4228:
4224:
4220:
4219:
4214:
4210:
4204:
4201:
4196:
4192:
4188:
4184:
4179:
4174:
4170:
4166:
4162:
4155:
4152:
4147:
4143:
4138:
4133:
4129:
4125:
4121:
4114:
4111:
4106:
4102:
4097:
4092:
4088:
4084:
4080:
4076:
4072:
4065:
4062:
4057:
4053:
4048:
4043:
4039:
4035:
4031:
4027:
4023:
4016:
4013:
4008:
4004:
4000:
3996:
3992:
3988:
3984:
3980:
3973:
3970:
3965:
3961:
3957:
3953:
3949:
3945:
3941:
3937:
3930:
3927:
3922:
3918:
3913:
3908:
3904:
3900:
3895:
3890:
3886:
3882:
3881:F1000Research
3878:
3871:
3868:
3856:
3852:
3848:
3842:
3839:
3834:
3830:
3824:
3822:
3818:
3813:
3809:
3804:
3799:
3795:
3791:
3787:
3783:
3779:
3772:
3769:
3764:
3760:
3756:
3752:
3748:
3744:
3737:
3730:
3728:
3724:
3719:
3715:
3711:
3707:
3703:
3699:
3692:
3689:
3683:
3678:
3674:
3670:
3666:
3659:
3657:
3653:
3642:on 2012-03-11
3638:
3634:
3630:
3626:
3622:
3618:
3614:
3607:
3600:
3597:
3592:
3588:
3583:
3578:
3574:
3570:
3566:
3562:
3558:
3551:
3548:
3543:
3539:
3535:
3531:
3527:
3523:
3519:
3515:
3511:
3507:
3500:
3497:
3492:
3486:
3482:
3478:
3474:
3467:
3464:
3453:on 2016-03-04
3452:
3448:
3444:
3440:
3436:
3432:
3428:
3424:
3417:
3414:
3409:
3405:
3400:
3395:
3391:
3387:
3383:
3379:
3375:
3368:
3366:
3364:
3360:
3355:
3351:
3347:
3343:
3336:
3333:
3328:
3324:
3319:
3314:
3310:
3306:
3302:
3298:
3294:
3287:
3284:
3279:
3275:
3271:
3267:
3263:
3259:
3255:
3251:
3244:
3241:
3236:
3232:
3227:
3222:
3218:
3214:
3210:
3206:
3202:
3195:
3192:
3187:
3180:
3177:
3173:(2): 387β400.
3172:
3168:
3164:
3157:
3154:
3149:
3145:
3141:
3137:
3133:
3129:
3125:
3121:
3120:
3115:
3108:
3105:
3100:
3096:
3092:
3088:
3084:
3080:
3076:
3072:
3065:
3062:
3057:
3053:
3049:
3045:
3041:
3037:
3033:
3029:
3022:
3019:
3014:
3010:
3006:
3002:
2998:
2994:
2990:
2986:
2985:
2977:
2974:
2962:
2961:New Scientist
2958:
2952:
2949:
2944:
2940:
2936:
2932:
2928:
2924:
2920:
2916:
2915:
2907:
2904:
2900:
2894:
2891:
2886:
2882:
2878:
2874:
2873:
2865:
2862:
2857:
2853:
2849:
2845:
2841:
2837:
2829:
2826:
2821:
2817:
2813:
2809:
2805:
2801:
2794:
2791:
2786:
2779:
2776:
2771:
2767:
2763:
2759:
2755:
2751:
2744:
2741:
2736:
2732:
2728:
2724:
2717:
2714:
2709:
2705:
2701:
2697:
2693:
2689:
2685:
2681:
2674:
2672:
2668:
2663:
2659:
2654:
2649:
2644:
2639:
2635:
2631:
2628:(6): e21086.
2627:
2623:
2622:
2617:
2610:
2607:
2602:
2598:
2593:
2588:
2583:
2578:
2574:
2570:
2569:
2564:
2557:
2554:
2549:
2545:
2541:
2537:
2533:
2529:
2525:
2521:
2514:
2511:
2506:
2502:
2495:
2492:
2487:
2483:
2479:
2475:
2468:
2465:
2460:
2459:Belknap Press
2456:
2449:
2446:
2441:
2437:
2433:
2429:
2425:
2421:
2417:
2413:
2412:
2407:
2400:
2398:
2396:
2392:
2387:
2383:
2379:
2375:
2371:
2367:
2363:
2359:
2355:
2351:
2344:
2342:
2340:
2336:
2330:
2325:
2321:
2317:
2314:(11): 27β54.
2313:
2309:
2305:
2298:
2296:
2292:
2287:
2283:
2278:
2273:
2268:
2263:
2259:
2255:
2251:
2244:
2241:
2229:
2225:
2221:
2217:
2213:
2209:
2205:
2201:
2197:
2193:
2190:(9): e03373.
2189:
2185:
2184:
2179:
2172:
2170:
2166:
2154:
2153:
2148:
2141:
2139:
2135:
2130:
2126:
2121:
2116:
2111:
2106:
2102:
2098:
2094:
2090:
2089:
2084:
2080:
2074:
2072:
2070:
2066:
2061:
2059:9780674454903
2055:
2051:
2046:
2045:
2039:
2033:
2030:
2026:(4): 215β216.
2025:
2021:
2014:
2012:
2008:
2003:
1999:
1995:
1991:
1990:
1982:
1980:
1976:
1971:
1967:
1962:
1957:
1953:
1949:
1945:
1941:
1937:
1933:
1932:
1927:
1920:
1917:
1913:(3): 375β393.
1912:
1908:
1904:
1900:
1894:
1892:
1888:
1883:
1879:
1875:
1871:
1867:
1861:
1859:
1857:
1855:
1853:
1849:
1843:
1838:
1835:
1831:
1828:
1825:
1822:
1821:
1820:
1817:
1815:
1812:
1810:
1807:
1805:
1802:
1800:
1797:
1795:
1792:
1791:
1786:
1784:
1782:
1781:
1776:
1772:
1771:
1766:
1762:
1758:
1757:
1752:
1745:
1743:
1741:
1737:
1731:
1729:
1723:
1721:
1717:
1712:
1711:
1702:
1700:
1698:
1694:
1690:
1689:
1683:
1681:
1677:
1676:
1670:
1668:
1664:
1663:
1658:
1653:
1652:
1643:
1638:
1634:
1632:
1627:
1625:
1621:
1617:
1613:
1609:
1608:
1602:
1595:
1590:
1588:
1584:
1577:
1575:
1573:
1572:kin selection
1568:
1564:
1558:
1550:
1548:
1546:
1542:
1541:H. rubicundus
1538:
1537:halictid bees
1532:
1530:
1521:
1519:
1515:
1512:
1504:
1502:
1498:
1496:
1492:
1487:
1484:According to
1479:
1475:
1471:
1466:
1461:
1460:Haplodiploidy
1457:
1449:
1444:
1442:
1439:
1435:
1434:W.D. Hamilton
1430:
1426:
1425:
1420:
1417:Prior to the
1412:
1405:
1404:
1396:
1395:
1392:
1390:
1386:
1385:
1381:
1380:Staghorn fern
1375:
1372:
1371:
1368:
1367:
1359:
1358:
1350:
1349:
1341:
1340:
1332:
1331:
1323:
1322:
1314:
1313:
1305:
1304:
1301:
1299:
1295:
1294:
1286:
1285:
1281:
1277:
1276:
1273:
1271:
1264:
1263:
1258:
1257:
1251:
1250:
1247:
1246:
1243:
1241:
1236:
1235:
1227:
1226:
1220:
1217:
1216:
1213:
1212:
1209:
1207:
1203:
1202:
1195:
1192:
1191:
1185:
1182:
1181:
1178:
1177:
1169:
1168:
1165:
1163:
1159:
1152:
1149:
1148:
1145:
1144:
1141:
1139:
1134:
1133:
1126:
1123:
1122:
1116:
1113:
1112:
1106:
1103:
1102:
1099:
1098:
1089:
1085:
1084:
1081:
1079:
1072:
1071:
1066:
1065:
1059:
1056:
1055:
1049:
1046:
1045:
1042:
1041:
1038:
1036:
1031:
1030:
1023:
1020:
1019:
1013:
1010:
1009:
1006:
1005:
1002:
1000:
996:
990:
987:
986:
980:
977:
976:
970:
967:
966:
962:
960:
955:
951:
947:
941:
933:
930:
922:
920:
918:
913:
908:
904:
903:Polypodiaceae
900:
899:
894:
886:
885:
879:
872:
870:
868:
864:
860:
856:
852:
851:kin selection
847:
845:
841:
836:
832:
829:or eusocial.
828:
822:
814:
812:
810:
806:
802:
798:
794:
790:
785:
783:
779:
775:
771:
767:
763:
759:
751:
746:
739:
737:
734:
730:
726:
725:
720:
712:
710:
707:
703:
700:
696:
692:
688:
684:
680:
676:
675:
670:
669:
664:
663:
658:
650:
648:
646:
642:
638:
634:
630:
629:
624:
620:
616:
615:
610:
606:
602:
594:
593:
588:
584:
580:
576:
569:
567:
565:
561:
557:
556:
548:
546:
544:
540:
537:, infraorder
536:
532:
525:
521:
514:
512:
510:
509:
504:
503:
497:
495:
490:
489:
484:
483:
477:
476:
475:Myrmecocystus
471:
470:honeypot ants
467:
458:
457:
456:Myrmecocystus
452:
448:
446:
445:
440:
439:
434:
433:
428:
424:
420:
416:
412:
408:
404:
400:
396:
392:
388:
385:) and wasps (
384:
380:
375:
373:
372:
367:
363:
359:
355:
351:
347:
343:
335:
334:
328:
321:
319:
315:
311:
303:
301:
299:
295:
291:
283:
281:
279:
271:
268:
265:
264:
263:
261:
254:
250:
243:
240:
237:
236:
235:
233:
229:
224:
222:
218:
214:
210:
206:
201:
197:
196:Suzanne Batra
189:
188:
183:
179:
178:Suzanne Batra
175:
168:
166:
164:
163:staghorn fern
160:
156:
152:
151:
146:
142:
138:
134:
129:
126:
122:
118:
114:
110:
106:
102:
98:
94:
90:
85:
83:
78:
74:
70:
66:
62:
58:
54:
50:
46:
39:
34:
30:
19:
7264:Sociobiology
7187:
7168:Allee effect
7142:Nanorobotics
7132:Ant robotics
7109:Vicsek model
6702:
6690:
6678:
6656:
6611:
6607:E. O. Wilson
6577:Noam Chomsky
6543:presociality
6530:
6435:Sociobiology
6372:
6365:
6358:
6351:
6344:
6303:E. O. Wilson
6253:Jane Goodall
6213:Donald Broom
6182:Zoosemiotics
6135:Sociobiology
5936:
5928:
5920:
5916:E. O. Wilson
5902:
5889:
5838:
5812:
5808:Thysanoptera
5798:
5640:Presociality
5620:
5555:
5549:
5543:
5534:
5530:
5520:
5469:
5463:
5457:
5412:
5406:
5366:
5362:
5356:
5323:
5317:
5311:
5294:
5288:
5230:
5226:
5222:
5154:
5148:
5138:
5129:
5125:
5119:
5076:
5070:
5060:
5017:
5011:
5001:
4974:
4970:
4960:
4935:
4931:
4927:
4891:
4887:
4877:
4860:
4854:
4850:
4844:
4828:(1): 12β22.
4825:
4819:
4813:
4805:
4764:
4758:
4733:(1): 17β25.
4730:
4724:
4711:
4668:
4664:
4650:
4607:
4603:
4593:
4560:
4556:
4547:
4504:
4500:
4490:
4460:(1): 17β52.
4457:
4453:
4432:
4426:
4417:
4408:
4385:
4376:
4368:
4363:
4350:
4344:
4301:
4297:
4287:
4268:
4264:
4255:
4222:
4216:
4203:
4168:
4164:
4154:
4130:(1): 72β78.
4127:
4123:
4113:
4078:
4074:
4064:
4029:
4025:
4015:
3982:
3978:
3972:
3939:
3936:Human Nature
3935:
3929:
3884:
3880:
3870:
3858:. Retrieved
3841:
3832:
3785:
3781:
3771:
3746:
3742:
3701:
3697:
3691:
3672:
3668:
3644:. Retrieved
3637:the original
3616:
3612:
3599:
3564:
3560:
3550:
3509:
3505:
3499:
3472:
3466:
3455:. Retrieved
3451:the original
3430:
3426:
3416:
3381:
3377:
3345:
3341:
3335:
3300:
3296:
3286:
3253:
3249:
3243:
3208:
3204:
3194:
3185:
3179:
3170:
3166:
3156:
3123:
3117:
3113:
3107:
3077:(1): 27β79.
3074:
3070:
3064:
3031:
3027:
3021:
2988:
2982:
2976:
2965:. Retrieved
2963:. 9 May 1992
2951:
2935:10.2307/4967
2918:
2912:
2906:
2893:
2876:
2870:
2864:
2839:
2835:
2828:
2803:
2799:
2793:
2784:
2778:
2753:
2749:
2743:
2726:
2722:
2716:
2691:
2687:
2683:
2679:
2625:
2619:
2609:
2572:
2566:
2556:
2523:
2519:
2513:
2504:
2500:
2494:
2477:
2473:
2467:
2454:
2448:
2418:(2): 86β87.
2415:
2409:
2405:
2353:
2349:
2311:
2307:
2257:
2253:
2243:
2231:. Retrieved
2187:
2181:
2156:. Retrieved
2150:
2092:
2086:
2043:
2032:
2023:
2019:
1993:
1987:
1935:
1929:
1919:
1910:
1906:
1873:
1869:
1778:
1768:
1764:
1761:ancient Rome
1754:
1749:
1739:
1732:
1724:
1708:
1706:
1686:
1684:
1673:
1671:
1660:
1649:
1647:
1628:
1605:
1599:
1585:
1581:
1567:E. O. Wilson
1560:
1545:L. calceatum
1544:
1540:
1533:
1525:
1516:
1508:
1499:
1495:haplodiploid
1490:
1483:
1470:haplodiploid
1422:
1416:
1382:
1379:
1378:
1291:
1289:
1267:
1260:
1232:
1230:
1199:
1198:
1155:
1130:
1129:
1125:Thysanoptera
1115:Paraneoptera
1075:
1068:
1027:
1026:
993:
943:
911:
896:
890:
882:
855:alloparental
848:
824:
808:
800:
786:
773:
765:
755:
732:
728:
722:
716:
708:
704:
698:
694:
690:
686:
682:
678:
672:
666:
660:
654:
633:Thysanoptera
626:
612:
598:
590:
578:
553:
552:
529:
506:
500:
498:
493:
486:
480:
473:
462:
454:
442:
436:
430:
376:
369:
339:
331:
317:
287:
275:
260:E. O. Wilson
258:
225:
193:
190:, pictured.
185:
155:E. O. Wilson
148:
125:reproductive
86:
77:reproductive
52:
44:
43:
29:
7188:Eusociality
7137:Microbotics
7007:butterflies
6978:sardine run
6912:altitudinal
6834:pack hunter
6652:Steven Rose
6531:eusociality
6243:Dian Fossey
6208:Marc Bekoff
6196:Ethologists
5814:Kladothrips
5750:Crabronidae
5735:Hymenoptera
5648:Agriculture
5621:Eusociality
5297:: 319β343.
4894:: 419β426.
2592:2434/801122
1996:: 299β342.
1876:: 109β115.
1736:royal jelly
1219:Hymenoptera
1132:Kladothrips
912:Platycerium
662:Synalphaeus
628:Kladothrips
543:pluripotent
515:In termites
438:Apis florea
387:Crabronidae
342:Hymenoptera
253:Weaver ants
205:oviposition
105:Hymenoptera
93:crustaceans
55:"good" and
45:Eusociality
18:Eusocialism
7248:Categories
7102:clustering
6995:philopatry
6973:salmon run
6968:Lessepsian
6642:Leon Kamin
6565:Supporters
6556:Polyethism
6145:Structures
6140:Stereotypy
5869:Coalescent
5857:In culture
5834:Coleoptera
5800:Synalpheus
5794:Crustacean
5755:Halictidae
5683:Polyethism
5495:2123/10940
5132:: 301β319.
4971:Apidologie
4726:BioScience
4625:10023/5520
4435:(1): 1β16.
3669:BioScience
3646:2011-04-04
3457:2007-11-30
3188:: 217β254.
3114:Synalpheus
2967:2010-10-31
2729:(2): 143.
2526:: 97β104.
2507:: 301β319.
1844:References
1756:Coalescent
1601:Pheromones
1596:Pheromones
1476:males and
1278:(100
1194:Coleoptera
1105:Eumetabola
1086:(150
1029:Synalpheus
1012:Arthropoda
969:Eukaryotes
768:) and the
758:Phiomorpha
733:H. pumilio
719:trematodes
668:S. regalis
657:crustacean
564:Coleoptera
549:In beetles
494:S. postica
427:pheromones
423:Euglossini
415:Meliponini
399:Polistinae
395:Formicidae
383:Halictidae
340:The order
308:See also:
304:In insects
290:arthropods
217:cell types
147:, such as
7223:Stigmergy
7203:Mutualism
6863:bait ball
6630:Opponents
6536:evolution
6526:Sociality
6514:in humans
6374:Behaviour
6317:Societies
6155:Honeycomb
5827:Aphididae
5822:Hemiptera
5760:Honey bee
5698:Thelytoky
5653:Gamergate
5126:Naturalia
4993:0044-8435
4977:: 13β30.
4507:: 15920.
4318:1559-2324
4239:0003-3472
3903:2046-1402
3718:1088-8683
3119:Evolution
2842:: 21β40.
2684:A. florea
2548:151303496
2501:Naturalia
2228:234496454
2212:0012-9658
1952:0066-4197
1819:Stigmergy
1680:pheromone
1642:fire ants
1612:aliphatic
1259:all
1151:Hemiptera
1067:all
1058:Blattodea
995:Mole-rats
940:Sociality
923:Evolution
893:epiphytic
873:In plants
827:prosocial
815:In humans
789:Carnivora
748:Model of
695:S. duffyi
687:S. chacei
619:Hemiptera
607:-forming
585:-forming
535:Blattodea
330:Swarming
284:Diversity
278:chordates
226:In 1969,
97:trematoda
69:offspring
61:sociality
38:honeybees
7152:Symbrion
7114:BIO-LGCA
6917:tracking
6846:ant mill
6788:sort sol
6783:flocking
6747:Swarming
6680:Category
6464:Ethology
6393:Category
6338:Journals
6165:Instinct
6115:Learning
6110:Instinct
6085:Ethogram
6068:Grooming
5991:Branches
5984:Ethology
5930:The Ants
5848:Isoptera
5772:Mammalia
5765:Vespidae
5564:43489291
5504:21516106
5449:20498859
5408:PLOS ONE
5247:84103230
5189:29403074
5111:21430721
5052:20740005
4952:44934383
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4797:20388889
4789:18511689
4703:20740005
4634:24021396
4539:28643786
4336:34338155
4247:53203398
4195:49380840
4187:29933836
4146:21185192
4105:30116921
4056:25673684
4007:37985774
3999:24347503
3964:44341304
3956:24091924
3921:27158472
3833:Prospect
3812:29072480
3763:16701397
3633:16701397
3591:25631992
3447:35627708
3408:39042696
3399:11295071
3327:44096968
3278:22548770
3235:19889706
3140:10937227
2820:36563930
2662:21695157
2621:PLOS ONE
2601:28343967
2540:31247426
2486:23736760
2455:The Ants
2440:35534268
2386:14607482
2286:11782550
2220:33988245
2129:16157878
1970:30208294
1787:See also
1740:Polistes
1716:oviposit
1710:Polistes
1667:exocrine
1620:releaser
1616:aromatic
1614:and two
1574:theory.
1237:(wasps)
1234:Vespidae
1184:Metabola
1156:various
1070:Termites
1022:Decapoda
989:Chordata
979:Animalia
954:Jurassic
867:meerkats
835:hominins
797:meerkats
793:Primates
729:in vitro
691:S. riosi
625:such as
539:Isoptera
531:Termites
524:Termites
472:such as
403:Vespinae
391:Vespidae
200:Halictid
182:Halictid
139:and the
121:termites
7012:monarch
6941:flyways
6922:history
6773:Droving
6692:Commons
6095:Feeding
5787:Meerkat
5777:Blesmol
5512:2060453
5474:Bibcode
5440:2871793
5417:Bibcode
5383:7247032
5348:9388017
5328:Bibcode
5180:6482461
5159:Bibcode
5102:3836173
5081:Bibcode
5043:3279739
5022:Bibcode
4769:Bibcode
4760:Science
4694:3279739
4673:Bibcode
4642:6548485
4585:1108197
4565:Bibcode
4557:Science
4530:5490048
4509:Bibcode
4482:5875340
4462:Bibcode
4327:8525959
4096:6289914
4047:4345452
3912:4850877
3860:31 July
3803:5730496
3582:4344137
3542:4251872
3534:8602260
3514:Bibcode
3305:Bibcode
3258:Bibcode
3226:2842683
3148:1088840
3099:8991755
3091:8603120
3056:4242926
3036:Bibcode
3013:2224506
2993:Bibcode
2923:Bibcode
2856:6867760
2770:4448321
2708:8657703
2653:3113908
2630:Bibcode
2420:Bibcode
2378:8029243
2358:Bibcode
2192:Bibcode
2183:Ecology
2120:1224642
2097:Bibcode
1961:6445553
1720:oophagy
1478:diploid
1474:haploid
1472:, with
1413:Paradox
1374:Plantae
1048:Insecta
631:(order
621:), and
617:(order
533:(order
419:Bombini
366:sterile
296:. Some
294:mammals
232:Apoidea
169:History
145:shrimps
143:. Some
133:rodents
101:mammals
89:insects
6985:Homing
6807:Locust
6442:Topics
5745:Apidae
5728:Groups
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5465:Nature
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1657:drones
1624:primer
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1158:Aphids
959:clades
907:fronds
865:, and
803:) and
778:inbred
697:, and
659:genus
623:thrips
592:Acacia
587:thrips
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379:Apidae
362:drones
358:queens
352:, and
213:tissue
159:humans
135:: the
115:, and
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7049:Boids
6990:natal
6778:Flock
6172:Swarm
6100:Hover
6053:Breed
5560:JSTOR
5508:S2CID
5344:S2CID
5243:S2CID
4948:S2CID
4793:S2CID
4638:S2CID
4243:S2CID
4191:S2CID
4003:S2CID
3960:S2CID
3739:(PDF)
3640:(PDF)
3609:(PDF)
3538:S2CID
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3323:S2CID
3144:S2CID
3095:S2CID
3052:S2CID
3009:S2CID
2939:JSTOR
2852:S2CID
2816:S2CID
2766:JSTOR
2704:S2CID
2544:S2CID
2482:JSTOR
2436:S2CID
2382:S2CID
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1826:(ACO)
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1665:have
1290:many
1231:many
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1032:spp.
609:aphid
599:Some
411:Apini
354:wasps
298:ferns
219:in a
128:males
117:wasps
65:brood
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6829:Pack
6795:Herd
6160:Nest
6150:Hive
5941:1994
5933:1990
5925:1975
5905:2000
5892:1927
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350:bees
346:ants
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