Eusociality

Eusociality (Greek εὖ eu "good" and social) is

the highest level of organization of sociality. It is
defined by the following characteristics:
cooperative brood care (including care of
offspring from other individuals), 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, sometimes called castes.
Eusociality is distinguished from all other social
systems because individuals of at least one caste
usually lose the ability to perform behaviors Co-operative brood rearing, seen here in honeybees, is
characteristic of individuals in another caste. a condition of eusociality.
Eusocial colonies can be viewed as
superorganisms.

Eusociality has evolved among the insects, crustaceans, trematoda and mammals. It is most widespread in
the Hymenoptera (ants, bees, and wasps) and in Blattodea (termites). A colony has caste differences:
queens and reproductive males 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 exert dominance by fighting. There are two eusocial rodents: the naked mole-rat and the
Damaraland mole-rat. Some shrimps, such as Synalpheus regalis, are eusocial. E. O. Wilson and others
have claimed that humans have evolved a weak form of eusociality. It has been suggested that the
colonial and epiphytic staghorn fern, too, may make use of a primitively eusocial division of labor.

History
The term "eusocial" was introduced in 1966 by Suzanne Batra, who used it to describe nesting behavior
in Halictid bees, on a scale of subsocial/solitary, colonial/communal, semisocial, and eusocial, where a
colony is started by a single individual.[1][2] 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,
oviposition) 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 superorganisms, with
individual castes being analogous to different tissue or cell types in a multicellular organism; 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.[3]
In 1969, Charles D. Michener[4] further expanded Batra's classification
with his comparative study of social behavior in bees. He observed
multiple species of bees (Apoidea) 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:[1]

1. Egg-layers and worker-like individuals among adult females

(division of labor)
2. The overlap of generations (mother and adult offspring)
Suzanne Batra introduced
3. Cooperative work on the cells of the bees' honeycomb
the term "eusocial"[1] after
E. O. Wilson extended the terminology to include other social insects, studying nesting in Halictid
such as ants, wasps, and termites. Originally, it was defined to include bees including Halictus
organisms (only invertebrates) that had the following three latisignatus,[2] pictured.
features:[1][5][6][7]

1. Reproductive division of labor (with or without sterile castes)

2. Overlapping generations
3. Cooperative care of young
Eusociality was then discovered in a group of chordates, 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 Weaver ants, here
sterile and provides support for the reproductive caste.[1][7] collaborating to pull nest
leaves together, can be
considered eusocial, as
Diversity they have a permanent
division of labor.

Most eusocial societies exist in arthropods, while a few are found in

mammals. Some ferns may exhibit a primitive form of eusocial
behavior.[8][9]

In insects
Eusociality has evolved multiple times in different insect orders, including hymenopterans,[10]
termites,[11] thrips,[12] aphids,[12] and beetles.[13]

In hymenoptera
The order Hymenoptera contains the largest group of eusocial insects, including ants, bees, and wasps—
divided into castes: reproductive queens, drones, more or less sterile workers, and sometimes also
soldiers that perform specialized tasks.[14] In the well-studied social wasp Polistes versicolor,[15]
 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.[16] Eusocial species with a sterile caste are
sometimes called hypersocial.[17]
Swarming Iridomyrmex purpureus While only a moderate percentage of species in bees (families
ants. The young queens are black,
Apidae and Halictidae) and wasps (Crabronidae and Vespidae) are
winged, and much larger than the
wingless workers.
eusocial, nearly all species of ants (Formicidae) are eusocial.[18]
Some major lineages of wasps are mostly or entirely eusocial,
including the subfamilies Polistinae and Vespinae. The corbiculate
bees (subfamily Apinae of family Apidae) contain four tribes of varying degrees of sociality: the highly
eusocial Apini (honey bees) and Meliponini (stingless bees), primitively eusocial Bombini (bumble bees),
and the mostly solitary or weakly social Euglossini (orchid bees).[19] Eusociality in these families is
sometimes managed by a set of pheromones that alter the behavior of specific castes in the colony. These
pheromones may act across different species, as observed in Apis andreniformis (black dwarf honey bee),
where worker bees responded to queen pheromone from the related Apis florea (red dwarf honey bee).[20]
Pheromones are sometimes used in these castes to assist with foraging. Workers of the Australian
stingless bee Tetragonula carbonaria, for instance, mark food sources with a pheromone, helping their
nest mates to find the food.[21]

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
self-sacrificing behavior. For example, members of the sterile
caste of the honeypot ants such as Myrmecocystus 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.[22] Not all social insects have distinct Myrmecocystus honeypot ants,
morphological differences between castes. For example, in the showing the repletes or plerergates,
Neotropical social wasp Synoeca surinama, caste ranks are their abdomens swollen to store
determined by social displays in the developing brood. [23] These honey (top), with ordinary workers
castes are sometimes further specialized in their behavior based on (bottom)
age, as in Scaptotrigona postica workers. Between approximately
0–40 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,
S. postica workers move outside the nest for colony defense and foraging.[24]
In Lasioglossum aeneiventre, 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.[25] In similar species with only one queen, such as Lasioglossum malachurum in
Europe, the degree of eusociality depends on the clime in which the species is found.[26]

In termites
Termites (order Blattodea, infraorder Isoptera) 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;[27] 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 pluripotent larvae
produced by the reproductive caste.[11] Some soldiers have jaws so enlarged
(specialized for defense and attack) that they are unable to feed themselves and
must be fed by workers.[28]
Termites live in large
In beetles nests, with queen,
king, soldier (red
Austroplatypus incompertus is a species of ambrosia beetle native to Australia,
heads), and worker
and is the first beetle (order Coleoptera) to be recognized as eusocial.[29][13] This (pale heads) castes.
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.[13]

In gall-inducing insects
Some gall-inducing insects, including the gall-forming aphid, Pemphigus
spyrothecae (order Hemiptera), and thrips such as Kladothrips (order
Thysanoptera), are described as eusocial.[12][30] These species have very
high relatedness among individuals due to their asexual reproduction
(sterile soldier castes being clones produced by parthenogenesis), 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, kleptoparasites, and competitors. In these groups,
Kladothrips rugosa, a gall-
eusociality is produced by both high relatedness and by living in a forming thrips (larva on left,
restricted, shared area.[31][32] adult on right) with galls
(centre) on Acacia leaves.
Its soldier caste defends the
In crustaceans colony in its gall fortress.
Eusociality has evolved in three different lineages in the colonial
crustacean genus Synalphaeus. S. regalis, S. microneptunus, S. filidigitus,
S. elizabethae, S. chacei, S. riosi, S. duffyi, and S. cayoneptunus 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.[33] They live eusocially with a single breeding female, and a large number of male defenders
armed with enlarged snapping claws. There is a single shared living space for the colony members, and
the non-breeding members act to defend it.[34]
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 system
(soldier caste); soldiers promote the fitness of the whole nest by ensuring safety and reproduction of the
queen.[35]

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.[36][37][38]

In trematodes
The trematodes are a class of parasitic flatworm, also known as flukes. One species, Haplorchis pumilio,
has evolved eusociality involving a colony creating a class of sterile soldiers.[39] 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.[40] 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.[40] Soldiers are readily distinguished from the immature and mature reproductive worms. Soldiers
are more aggressive than reproductives, attacking heterospecific trematodes that infect their host in vitro.
H. pumilio 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
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".[40]

In nonhuman mammals
Among mammals, two species in the rodent group
Phiomorpha are eusocial, the naked mole-rat
(Heterocephalus glaber) and the Damaraland mole-rat
(Fukomys damarensis), both of which are highly
inbred.[41] 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 avoid
inbreeding, mole rats sometimes outbreed and
Model of naked mole-rat burrow with soldiers,
establish new colonies when resources are
workers, and queen, a social structure similar to
sufficient.[42] Most of the individuals cooperatively the castes of the eusocial insects
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.[43]
Some mammals in the Carnivora and Primates have eusocial tendencies, especially meerkats (Suricata
suricatta) and dwarf mongooses (Helogale parvula). 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.[44]

In humans
Scientists have debated whether humans are prosocial or eusocial.[45] Edward O. Wilson called humans
eusocial apes, arguing for similarities to ants, and observing that early hominins cooperated to rear their
children while other members of the same group hunted and foraged.[46] Wilson and others argued that
through cooperation and teamwork, ants and humans form superorganisms.[47][48][49] Wilson's claims
were vigorously rejected by critics of group selection theory, which grounded Wilson's
argument,[46][50][51] and because human reproductive labor is not divided between castes.[50]

Though controversial,[52] it has been suggested that male homosexuality[53] and female menopause[54]
could have evolved through kin selection.[55][56] This would mean that humans sometimes exhibit a type
of alloparental behavior known as "helpers at the nest", with juveniles and sexually mature adolescents
helping their parents raise subsequent broods, as in some birds,[57] some non-eusocial bees, and
meerkats.[58] These species are not eusocial: they do not have castes, and helpers reproduce on their own
if given the opportunity.[59][48][60]

In plants
One plant, the epiphytic staghorn fern, Platycerium bifurcatum
(Polypodiaceae), 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 fronds 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 Platycerium 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 specialized for tasks, a division of The staghorn fern
labor.[8][9][61] Platycerium bifurcatum may
display a simple form of
eusociality.

Evolution

Phylogenetic distribution
Eusociality is a rare but widespread phenomenon in species in at least seven orders in the animal
kingdom, as shown in the phylogenetic tree (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
Jurassic period (~150 million years ago).[62] 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 (clades) is not known. The major eusocial groups are
shown in boldface in the phylogenetic tree.

⁠Chordata Mole-rats
⁠

⁠Decapoda Synalpheus spp.

⁠

⁠Blattodea ⁠all Termites

⁠ ⁠(150 mya)

⁠Thysanoptera Kladothrips
⁠
⁠Paraneoptera
⁠

⁠Hemiptera various Aph

⁠
⁠Animalia
Austroplaty
⁠Arthropoda
⁠Eukaryotes ⁠ ⁠Coleoptera
⁠ ⁠
⁠Insecta
⁠
⁠Eumetabola
many
⁠
⁠
⁠

⁠Metabola
⁠
⁠Hymenoptera
⁠ ll Ants
a
⁠
⁠(100 mya)
⁠
⁠
⁠
⁠

⁠Plantae
⁠
Paradox
Prior to the gene-centered view of evolution, 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 On the Origin of Species, Darwin 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".[63] Darwin anticipated that a possible resolution to the paradox might lie in the close
family relationship, which W.D. Hamilton quantified a century later with his 1964 inclusive fitness
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.[64]

Inclusive fitness and haplodiploidy

Argument that haplodiploidy favors eusociality

According to inclusive fitness
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 the fraction of
genes that they share, i.e. when
Cost < relatedness * Benefit. W.
D. Hamilton suggested in 1964
that eusociality could evolve
more easily among haplodiploid Honey bees are haplodiploid, with haploid males and diploid females. It
species such as Hymenoptera, has been suggested that this organisation favours eusociality, but
haplodiploidy is neither necessary nor sufficient for eusociality to emerge.
because of their unusual
relatedness structure.[65][66][67]

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 to what W.
D. Hamilton first termed "supersisters", more closely related to their sisters than they would be to their
own offspring.[67] 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.[68]

Argument that haplodiploidy does not favor eusociality

Against the supposed benefits of haplodiploidy for eusociality, Robert Trivers notes that while females
share 75% of genes with their sisters in haplodiploid populations, they only share 25% of their genes with
their brothers.[69] Accordingly, the average relatedness of an individual to their sibling is 50%. Therefore,
helping behavior is only advantageous if it is biased to 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.[70]

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 association
between haplodiploidy and eusociality is below statistical significance.[71] Haplodiploidy is thus neither
necessary nor sufficient for eusociality to emerge.[72] Relatedness does still play a part, as monogamy
(queens mating singly) is the ancestral state for all eusocial species so far investigated.[73] 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.[73]

Evolutionary ecology
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.[7] The importance of
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.[74]
Conversely, female Damaraland mole-rats undergo hormonal changes that promote dispersal after periods
of high rainfall.[75]

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.[76] Similarly, social transitions within halictid bees, 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 H. rubicundus, 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.[77] In another species of
sweat bees, L. calceatum, 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.[78]

Multilevel selection
Once pre-adaptations such as group formation, nest building, high cost of dispersal, and morphological
variation are present, between-group competition has been suggested as a driver of the transition to
advanced eusociality. M. A. Nowak, C. E. Tarnita, and E. O. Wilson 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.[79] Multilevel
selection has been heavily criticized for its conflict with the kin selection theory.[80]

Reversal to solitarity
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 bees was
followed by at least one reversal to solitarity, giving a total of at least nine reversals.[4][5] 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.[77] 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.[81]

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 the
honeybee from reverting to the solitary state.[20]

Physiology and development

Pheromones
Pheromones 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.[82] The best-studied queen pheromone system in social insects is that
of the honey bee Apis mellifera. Queen mandibular glands produce a mixture of five compounds, three
aliphatic and two aromatic, which control workers.[83] Mandibular gland extracts inhibit workers from
constructing queen cells, which can delay the hormonally based behavioral development of workers and
suppress their ovarian development.[84][83] Both behavioral effects mediated by the nervous system often
leading to recognition of queens (releaser) and physiological effects on the reproductive and endocrine
system (primer) 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.[84]
The levels of two of the aliphatic compounds increase rapidly in virgin queens within the first week after
emergence from the pupa, consistent with their roles as sex attractants during the mating flight.[83] Once a
queen is mated and begins laying eggs, she starts producing the full blend of compounds.[83] In several
ant species, reproductive activity is associated with pheromone production by queens.[83] Mated egg-
laying queens are attractive to workers, whereas young winged virgin queens elicit little or no
response.[83]

Among ants, the queen

pheromone system of the fire
ant Solenopsis invicta 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 Physiology of eusociality in fire ants: three queen pheromones help to
initiate reproductive create and maintain the eusocial state of the colony. Loss of a primer
development in new winged pheromone triggers the development of replacement queens (dashed
females, called female lines).[83][84]
sexuals.[83] 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.[83][84] These pheromones maintain the eusocial phenotype, with one
queen supported by sterile workers and sexually active males (drones). 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.[83]
Similarly, queen weaver ants Oecophylla longinoda have exocrine glands that produce pheromones
which prevent workers from laying reproductive eggs.[84]

Similar mechanisms exist in the eusocial wasp Vespula vulgaris. 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 pheromone from the queen's body alerts
those workers of her dominance.[85]

The mode of action of inhibitory pheromones which prevent the development of eggs in workers has been
demonstrated in the bumble bee Bombus terrestris.[84] The pheromones suppress activity of the endocrine
gland, the corpus allatum, stopping it from secreting juvenile hormone.[86] 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.[84]

Other mechanisms
A variety of other mechanisms give queens of different species of social insects a measure of
reproductive control over their nest mates. In many Polistes 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 rank of subordinates is correlated with the degree
of ovarian development.[84] Workers do not oviposit when queens are present, for a variety of reasons:
colonies tend to be small enough that queens can effectively dominate workers; queens practice selective
oophagy; the flow of nutrients favors queen over workers; and queens rapidly lay eggs in new or vacated
cells.[84]

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 and
maintenance.[84] Being nudged by the queen may help to inhibit ovarian development; in addition, the
queen eats any eggs laid by workers.[84] Furthermore, temporally discrete production of workers and
gynes (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.[87] In
some cases, for example in bumblebees, queen control weakens late in the season, and the ovaries of
workers develop.[84] 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.[84]

In highly eusocial wasps (where castes are morphologically dissimilar), both the quantity and quality of
food are important for caste differentiation.[84] Recent studies in wasps suggest that differential larval
nourishment may be the environmental trigger for larval divergence into workers or gynes.[87] All honey
bee larvae are initially fed with royal jelly, 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.[88] The
differential expression in Polistes 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.[87]

In popular culture
Stephen Baxter's 2003 science fiction novel Coalescent imagines a human eusocial organisation founded
in ancient Rome, in which most individuals are subject to reproductive repression.[89] Harold Fromm,
reviewing Groping for Groups by E. O. Wilson and others in The Hudson Review, 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 Huxley's [1932 novel] Brave New World.[90]

See also
Dense heterarchy
 Evolutionarily stable strategy
International Union for the Study of Social Insects
Patterns of self-organization in ants
Reciprocity (social psychology)
Stigmergy
Ant colony optimization (ACO)
Bee colony optimization
Task allocation and partitioning of social insects

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External links
International Union for the Study of Social Insects (http://www.iussi.org/)
Eusociality in naked mole-rats (http://academic.reed.edu/biology/professors/srenn/pages/tea
ching/web_2007/molerats_lb_jg/index.html)

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