THEORIES ABOUT

EVOLUTION OF SOCIALITY
IN INSECTS AND FACTORS
ALI HASSAN
2018-AG-6628
AQUATIC AND SOCIAL INSECTS
ENT-514
Department of entomology
University of agriculture faisalabad
Theories related to evolution
Kin selection
 Kin selection has been used to explain evolution of many altruistic traits found in those social insect castes which
have reduced reproductivity.
Kin selection is most probable among organisms living in family groups. It is commonly assumed, therefore, that:
I. The most primitive social insects, like the more specialized ones, live in mother-daughter (eusocial) colonies,
II. Colonies of individuals of the same generation and showing castes (semisocial colonies) are abnormal or originate
only in inbreeding populations, and
III. Transfer of individuals among colonies must be selected against, except in inbreeding populations.
 Kin selection is selection among kin- ships or families rather than among individuals. The selection operates
directly upon individuals without reproductivity or with reduced reproductivity (workers or soldiers).
 But since these individuals contribute to the welfare of the whole family, i.e., the colony, their success influences
the reproductivity of the queen who produces the sexual forms of the next generation
Hamilton (1964

 Hamilton (1964; in press) has treated such kin selection in detail,

placing emphasis on the proportion of genes identical by descent
(among siblings, between parents and offspring, or between other
relatives) necessary for altruistic traits to arise.
 Altruistic traits arise only through kin selection, i.e., they arise when
the advantages of altruistic behavior and also the level of
relationship among the individuals, together representing inclusive
fitness, are high enough.
 One can also say that kin selection is selection acting on inclusive
fitness, although, of course, the latter is always composed of diverse
properties upon which diverse selective forces (jointly kin selection)
must be operating.
William Hamilton's concept of inclusive fitness

 "I had realized from experience that university people sometimes don't react well to common sense
and in any case most of them listen to it harder if you first intimidate them with equations."
 Hamilton Traditional "direct" fitness is counted as the number of offspring produced. Inclusive fitness
means that you can still have some fitness if you yourself produce no offspring-by considering other
individuals that share genes with you.
 For example, you share 50% of genes with your children-hence having two children will mean, on
average, that you have produced one full copy of your genome. (Hence if you commit altruistic suicide
to save more than 2 of your children, you have won!)
 Any full sibling (brother or sister) shares 50% of your genes, and their children share 25% of their
genes with you.
 Thus, being an uncle or aunt to 4 children is as good as having 2 children of your own!
 You can increase your own fitness by helping a close relative bring up children!
Trivers (1971)

 Trivers (1971) uses the term "reciprocal altruism" for behavior such that animal Number 1, at risk
to its own productivity, enhances that of Number 2, but such that later the positions may be
reversed, so that Number 1 will benefit at risk to Number 2.
 Depending on the levels of benefits and risks involved, such behavior can be advantageous to both
individuals and favored by selection at the individual level.
 The word "altruism" is in a sense a misnomer. Activity that promotes the perpetuation of an
individual's own genes directly is called "selfish" but promotion of the same genes via another
individual is called "altruistic."
 Altruism in the strict sense of the promotion of others' welfare, without reward, is really unknown
(except as accidents or as culturally conditioned phenomena in man).
 In the long run, reciprocally altruistic behavior is a class of mutualistic social behavior
Michener (1969)

 A colony consists of the interacting individuals of a species in one nest (young ones sealed in cells are
excluded). A communal colony consists of individuals (of a single generation), with no division of labor.
 A semisocial colony consists of adult individuals of a single generation among which there is division of
labor, a queen and workers, or at least an approach to such castes, because of considerable variation in
productivity and foraging activity.
 A eusocial colony consists of adults of two generations, the queen being the mother of the workers. In
primitively eusocial colonies, queens and workers are morphologically about alike;the potential queens
have the ability to live alone.
 In highly eusocial colonies queens and workers are quite different; the queens often lack the ability to live
alone or if they do so, they commonly do not forage (as in colony establishment of most ants), and the
colonies are long-lived.
 A subsocial colony consists of a mother and her immature offspring (nymphs, larvae), the latter being
protected and usually fed by the mother, and in frequent contact with her. The colony breaks up be- fore
the young become adults, and there is no colony of adult.
Crozier (1970


Male haploidy must initially have resulted in a reduction of genetic variability. This statement applies especially to
highly deleterious alleles. Balanced polymorphisms may be as common in haplo-diploid species as in fully diploid
ones. At any one fertilization, only three-fourths as many chromosomes are involved as in a fully diploid system.

Haploid male bee copulates with diploid female -> haploid sperm is stored in female bee's spermatheca.When
female"wants" to produce son, she lays an unfertilised (haploid) egg -> male offspring.

To produce female offspring, mother needs to add sperm to egg as it passes down oviduct. 
Thus haplo-diploidy implies that:

 It might be more beneficial for a worker to help raise more sisters (workers and new queens) than to reproduce on
her own 

This is because the average relationship between sisters is higher (0.75) than between workers and their (possible)
offspring (0.5)

 This might explain why the haplodiploid Hymenoptera have evolved eusocial systems more than any other insect
order
Male haploidy is related to many aspects of
hymenopteran evolution.

Among these is the origin and evolution of social behavior because of

 Partial immunity to inbreeding resulting from the loss of lethals, allowing
long-term aggregations and colonies and effective kin selection to arise,
 Sperm storage by females, absence of a king in social groups, and the control
of the sex of offspring, permitting the extra- ordinary sex ratios found in
highly social Hymenoptera,
 The importance of male- production by workers in primitively social groups
and some highly social ones, and
 The closer relation of sisters to one another than to their daughters, an
arrangement that encourages effective kin selection
Individual Selection

 There is considerable variation in reproductivity of females even in solitary forms;

 Highly productive individuals are likely to make their nests first, before the less productive ones, and thus
increase their advantage.
 The percentage of mortality among the offspring of all individuals in a colony is equal, and is lower than that
experienced when individuals are alone in their own nests.
 It is therefore safe to conclude that laying by workers has a reproductive function important enough for selection
to maintain the reproductive system and reproductive behavior.
 This fact does not mean that kin selection, producing altruism, has not been involved in the evolution of a worker
caste, but it does mean that some individual selection, leading to mutualistic behavior, has been involved.
 Presumably there has been an interaction between the two kinds of selection. The larger the productivity of the
joiner or worker, the less relevant is the coefficient of relationship to an understanding of the evolution of the
worker caste.
 If kin selection were highly pre- dominant over individual selection, one would expect the reproductive system
to be functionless and vestigial or absent in workers of the more highly social Hymenopter
MULTIPLE SELECTION

The following can be interpreted as counter- measures-devices that increase variability:

 Multiple insemination appears to occur in various groups (some ants, e.g., Atta, whose queens mate 3 to 8
times -Kerr, 1961; Apis; some other bees, etc.)
 multiple queens, occurring either simultaneously as in polygynous (pleiometrotic) forms (see Wilson, 1963b)
or successively (as in Apis)
 queen, worker, or male transference or joining
 Worker laying of male-producing eggs and sometimes of female eggs
 Worker mating (as in some ants and some primitively social bees)
 Common mating places where young gynes and males from different colonies in a considerable area
congregate (ants, honey bees, Polistes)
 Synchronization of mating for reproductives from different colonies in a large area. Most of these
mechanisms occur in other groups as well, but their frequency in the Hymenoptera is impressive and may be
related to maintenance of an adequate level of genetic variability.
Factors of evolution

1. Influence of Natural Enemies

2. Selection for Joining and Accepting Behavior
Patterns
3. Familial and Semicocial Evolution Of Sociality in
Hymenptea
4. Monogamy
5. Ecological factors
6. Genetic influences
Influence of Natural Enemies
 At least in halictine bees, a major selective factor favoring the establishment of colonies, irrespective of their mode
of origin, is almost surely parasite and predator pressure (Michener, 1958; Michener and Lange, 1958; Lin, 1964a).
 In ground-nesting groups like the Halictinae, this means pressure of the natural enemies in aggregations of nests;
scattered nests presumably experience less such pressure.
 Social vespids may never have passed through an aggregating phase, but they and their nests are large and
conspicuous, and again it seems possible that natural enemies were a major selective factor leading to the
establishment and enlargement of colonies.
 Improved control of other environmental factors in the nest and economy of labor in nest-making may also have
been involved.
 Pressure by parasites or predators probably varies enormously and is likely to become devastating in some
aggregations of nests. Michener (1966a) has seen an aggregation of Lasioglossum versatum decimated by
mutillids, some of them large, of L. zephyrum decimated by mutillids, by myrmosids, and by a combination of
mutillids and ant.
 Parasite and predator pressure is perhaps unusually high for aggregating species of halictines because of their
round, perpendicular (hence conspicuous) nest entrances. Perhaps this is one reason for the frequent origin of
sociality in this group
Selection for Joining and
Accepting Behavior Patterns

 Irrespective of the evolutionary route, the evolution of joining and accepting behavior
and the beginning of semisocial or eusocial colonies can be accounted for if we make
the assumptions that
 (1) there is considerable variation in reproductivity of females even in solitary forms
 (2) highly productive individuals are likely to make their nests first, before the less
productive ones, and thus increase their advantage
 (3) the percentage of mortality among the offspring of all individuals in a colony is
equal, and is lower than that experienced when individuals are alone in their own
nests.
Familial and Semicocial Evolution
Of Sociality in Hymenptea
 Once a joining class exists and is advantageous, selection would probably favor
colonies that produce their own joiners.
 Ordinarily this means colonies that are eusocial. Such colonies would in that case be
surer of getting joiners than if the joiners came from other nests.
 A eusocial society might be established secondarily, superimposed on or derived
from a semisocial society, whether or not kin selection was a predominant factor in
establishment of the semisocial organization.
 After such eusocial family groups exist, evolution of special queen and worker
features, including reduction in worker productivity, becomes possible by kin
selection.
Monogamy

  Natural selection will favor cooperation in any situation where it is more efficient to raise siblings than
offspring, and this could start paving a path towards sociality. This higher efficiency becomes especially
pronounced after group living evolves.
 In many monogamous animals, an individual's death prompts its partner to look for a new mate, which
would affect relatedness and hinder the evolution of sociality: workers would be much more related to their
offspring than their siblings.
 However, many Hymenoptera have a form of lifetime monogamy in which the queen mates with a single
male, who then dies before colony establishment. This seems to be the ancestral state in all Hymenopteran
lineages that have evolved sociality.
  Most termites also have a mating system in which a reproductive female (the queen) commits to a single
male for life (the king), and this pattern seems to be ancestral in termites. Lastly, strict monogamy facilitated
sociality in the sponge-dwelling shrimp.
Ecological factors

 Ecological factors were also probably a precursor to sociality.

  For example, the sponge-dwelling shrimp depend upon the sponge's feeding current
for food, termites depend upon dead, decaying wood, and naked mole rats depend
upon tubers in the ground.
  Each of these resources has patchy distributions throughout the environments of
these animals.
 This means there is a high cost to dispersing (individual may not find another source
before it starves), and these resources must be defended for the group to survive.
 These requirements make it a necessity to have high social order for the survival of
the group.
Genetic influences

 Genetic constraints may have influenced the evolution of sociality.

 The genome structure of the order Hymenoptera has been found to have the
highest recombination rate of any other groups in Animalia.
 The eusocial genus Apis, the honeybees, have the highest recombination rate in higher
eukaryotes. Genes determining worker behavior and division of labor have been found in regions
of the Apis genome with the highest rates of recombination and molecular evolution. 
 These mechanisms are likely important to the evolution of sociality because high recombination
rates are associated with the creation of novel genes, upon which natural selection can act. This
could have been important in other eusocial genera.
 Biased gene conversion rates are also higher in eusocial species.This could increase genotypic
diversity, which could allow workers to meet the demands of a changing social structure more
easily.
  Another hypothesis is that the lower overall genetic diversity as sociality levels increase
throughout the family Apidaeis due to a decreased exposure to parasites and pathogens.
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