author={David C. Queller and Joan E. Strassmann},
Social insects so dominate many terrestrial habitats (Wilson 1990) that they can hardly escape the attention of biologists, but even if they were rare, they would still attract special interest because of the intricate cooperation within their societies. William Morton Wheeler (1911) described the social insect colony as an organism (or as a higher-level organism or superorganism) because of the degree to which individuals appear to operate as a unit that is dedicated to the perpetuation and… 
Insect societies as divided organisms: The complexities of purpose and cross-purpose
Although kin selection explains the extensive cooperation and common purpose of social insect colonies, it also predicts a certain amount of cross-purpose and conflict behavior.
Social insects, major evolutionary transitions and multilevel selection
The multilevel selection approach is presented, which promises to be a useful tool to study evolution at multiple selection levels and to quantify benefits and costs of cooperation, so that insect societies and all major evolutionary transitions alike are being recognised as more than the sum of their components.
Multilevel selection and social evolution of insect societies
Insect societies can be regarded as a level of selection with novelties that provide benefits beyond the scope of a solitary life, and relatedness is less important for the maintenance of insect societies, although it played a fundamental role in their evolution.
Unrelated helpers in a social insect
Microsatellite markers are used to reveal an unexpected and unique social system in what is probably the best-studied social wasp, Polistes dominulus, which is functionally unlike other social insects, but similar to certain vertebrate societies, in which the unrelated helpers gain through inheritance of a territory or a mate.
An ecological driver for the macroevolution of morphological polymorphism within colonial invertebrates
  • Carl Simpson
  • Biology
    Journal of experimental zoology. Part B, Molecular and developmental evolution
  • 2020
This paper proposes a mechanism that links the origin of novel body types to the evolution of life‐history strategies among species, and suggests that each new polymorph type that evolves permits more variation in colonial life‐histories to exist.
The Ecology of Altruism in a Clonal Insect
The ecological context of altruism in social aphids has been shown to be quite intricate since it is now clear that colony defense is not the only costly behavior that they perform: they also have vital roles in keeping the colony clean, migrating to new colonies, and repairing their nest.
Evolution of supercolonies: The Argentine ants of southern Europe
It is suggested that a “genetic cleansing” of recognition cues occurred after introduction of the Argentine ant, which resulted in the formation of two immense supercolonies, one of which effectively forms the largest cooperative unit ever recorded.
Relatedness and the fraternal major transitions.
  • D. Queller
  • Biology
    Philosophical transactions of the Royal Society of London. Series B, Biological sciences
  • 2000
The role of kinship is examined, focusing on the transitions to multicellularity in animals and to coloniality in insects, to show that kin selection based on high relatedness permitted cooperation and a reproductive division of labour.
It is found that native populations also form su- percolonies, and are effectively unicolonial, just as in introduced populations, and the relatedness between nestmates is not distinguishable from zero in these native range supercolonies.
Nonrelatives inherit colony resources in a primitive termite
Findings demonstrate how ecological factors could have promoted the evolution of eusociality by accelerating and enhancing direct fitness opportunities of helper offspring, rendering relatedness favoring kin selection less critical.


Commentary: Extrinsic versus Intrinsic Factors in the Evolution of Insect Sociality
Over the past few years, there has been a veritable flood of speculation regarding the evolution of social behavior, especially in Hymenoptera. A partial list of papers might include Lin and Michener
Haploidploidy and the evolution of the social insect.
Evidence is presented from 20 species that the ratio of investment in monogynous ants is, indeed, about 1 : 3, and this discovery is subject to a series of tests, which provide quantitative evidence in support of kinship theory, sex ratio theory, and the assumption that the offspring is capable of acting counter to its parents' best interests.
The Evolution of Eusociality
An evaluation of the various hypotheses proposed to explain the evolution of eusociality in Hymenoptera and Isoptera shows that genetic related­ ness between the workers and the reproductives they help raise may resolve the apparent paradox.
Evolution of eusociality: the advantage of assured fitness returns
It is shown that an `assured fitness returns' model provides a selective pressure for the evolution of worker behaviour which is at least about as strong as that of haplodiploidy, but free from such requirements of the latter as high levels of worker-brood genetic relatedness and ability of workers to manipulate brood sex ratios.
Developmental and Physiological Determinants of Caste in Social Hymenoptera: Evolutionary Implications
A developmental view of the evolution of societies is presented and two complementary questions are raised regarding the relationship between social structure and the factors that determine physical or physiological differences among females are raised.
A selfish strategy of social insect workers that promotes social cohesion
It is shown that when colonies have multiple queens born in the same nest, the selfish worker sex-ratio strategy has a paradoxical side-effect which strongly promotes social cohesion, and may be responsible for the maintenance of eusociality in this group.
Eusociality in a coral-reef shrimp
Ecological similarities among mole-rats, termites and these sponge-dwelling shrimp, all of which are diploid animals, strengthen arguments that eusociality is favoured by gradual metamorphosis, parental care, and occupation of protected, expansible niches.
Regulation of Reproduction in Eusocial Hymenoptera
An evolutionary framework focuses attention on the social factors that regulate reproduction, but to avoid separating them artificially from their ecological context, factors extrinsic to a colony must occasionally enter the discussion.
The evolution of eusociality: Reproductive head starts of workers.
  • D. Queller
  • Biology
    Proceedings of the National Academy of Sciences of the United States of America
  • 1989
In eusocial species, many individuals forego their personal reproduction to aid the reproduction of their mother or other relatives. Kin selection can favor such behavior for any positive degree of
Evolution of Colony Characteristics in Social Insects. I. Sex Allocation
  • P. Pamilo
  • Economics
    The American Naturalist
  • 1991
The inclusive-fitness model elaborated in this article can be applied to derive the expected allocation patterns of social insects, both as a function of the type of colony and of the amount of resources the colony can use in sexual production.