How is the Social and Mating Behaviour
of a Variety of Species related to their Ecology and Evolution?
The
evolutionary advance of sociality and social structure is an important
selective factor which affects an animal’s survival and reproductive success (Ilany & Akçay, 2016). Sociality plays an important
role within health and evolutionary fitness within an individual; affecting
pathogen transmission and increasing/decreasing particular social behaviours (Hinde,
1976). Social structures
within a population show the social bonds and patterns which occur between each
member as a group and as an individual (Wey,
Blumstein, Shen, & Jordán, 2008).
Mating systems are classed as the way
animal populations are structured in relation to reproductive/sexual behaviour
(Stutchbury, 1998). A mating system is
the way males or female’s pair when choosing a mate; males and females fluctuate
significantly in the investment they make to reproduce, and therefore tackle
mating with a range of different strategies (Klusmann,
2004). When choosing a mate
certain species have evolved species-typical strategies to take get the best
reproductive success; which has created a large diversity amongst species and
their mating patterns (McCracken & Bradbury,
1981)
Socioecology is the study of how
social structures are changed or influenced by the environment. It focuses on
males and female’s behavioural responses to regards to resources, distribution
and predation risk (Rubenstein, 1987).
Within this essay the species differences in grouping structure and
social relationship patterns will be discussed in termites, honey bees and ants;
three highly eusocial animals(Varley
& Klopfer, 1963).
Termite social system
Eastern subterranean termites (Reticulitermes
flavipes) eusocial animals
meaning they are at the highest level of animal sociality ((Howard, Jones, Mauldin, & Beal, 1982) their
colonies can range from 100,000
up to 1 million individuals composed of both males and females (Thorne, Traniello, Adams, &
Bulmer, 1999). Their colonies are classed as
“super-organisms” as the termites self-regulate the colony. Termites have a
caste system; which consist of, workers, soldiers and reproductive (Noirot, 1989).
Workers: make up the majority of colony.
Their responsibilities are foraging for food/water, feeding and grooming other termites,
caring for the queen and her eggs. Alongside constructing and repairing the
tunnels, workers are sterile and therefore unable to breed within the colony (Higashi, 2000).
Soldiers: primary function is to
defend the colony. Soldier termites are
incapable of feeding themselves and they reply upon the worker ants to provide
them with food that has been regurgitated (Mao, Henderson, Liu, &
Laine, 2005).
Reproductives:
essential part within the colony, they take part in a mating swarm once weather
conditions are acceptable. After which they will go and form new colonies, and
become the reproductive king and queen of that colony (Noirot, 1990).
Termite mating system
Termite kings and
queens are monogamous, while the lower termites take part in polygamy (Wu et al., 2014). The king’s role is to stay with the queen and
reproduce in order to form a successful colony (Weaver, 1971). The queen’s role
in the colony is egg production in order to grow the colony. She lays eggs
daily and reaches her peak production rate between seven and ten years old
(Weaver, 1971:Hartke & Baer, 2011).
Primary
reproductives: consist of queens, kings and alates. During nuptial flight, the
queen produces alates which leave the nest and new colonies. When forming new
colonies new nest sites will be located, the queen will then lay eggs to
produce workers, which will then care for her eggs alongside expanding the nest
(Reilly, 1987:Matsuura, 2010).
Secondary and
tertiary reproductives: the queen controls the size of the colony, therefore
preventing secondary and tertiary reproductives forming by pheromones (Howard and
Haverty, 1980). The queen will do this when the colony reaches a size the
queen is happy with, the king and queen can also produce a pheromone which
circulates the colony to stop other members reproducing (Thorne and
Noirot, 1982:Hartke and Baer, 2011)
Nuptial nights depend
on environmental conditions, such as rain fall or moisture. In some colonies
queens are able to switch their mating from mating by sexual reproduction to
asexual production, the queen can therefore produce replacement workers
parthogenetically (Williams,
1959:Henderson and Delaplane, 1994).
Honey bee social system
Honey bees (Apis mellifera) are eusocial bees, their
colonies will generally contain a single queen bee, a few thousand of drone
bees and tens of thousands of worker bees (Dolezal and
Toth, 2013).
Drones: the male bees otherwise known
as drones, their only purpose is to mate with the queen which then they
immediately die afterwards (Giray and Robinson, 1994:
Schulz, Sullivan and Robinson, 2002).
Workers: are female bees, due to a
pheromone released by the queen the workers are sterile and therefore unable to
reproduce. They are responsible for everything within the hive, cleaning,
collecting food, making new cells as well as being the main factor to if the
hive moves or swarms. Workers are also able to develop ovaries if the colony loses
their queen (Giray and Robinson, 1994: Schulz, Sullivan
and Robinson, 2002).
Queens: the main female reproductive, she takes flight
around the hive to male with the drones and can lay up to 10,000 eggs in one
day. (Giray and Robinson, 1994: Schulz, Sullivan and
Robinson, 2002).
Honey bee mating system
Polyandry is practiced by the queen,
as she has multiple sexual interactions with the males. Queens fly to sites
where thousands of males will be waiting of which she will mate with several
during flight (Pechacek et al., 2005). Males can
only mate seven to ten times during a mating flight, as their abdomen rips open
when their endophallus enters inside the queen and leads to death (Chiu and Kuo, 2010).There sole purpose is for mating
so even if the mating flight is survived they will be ejected from their
previous nest as their purpose has been served.
Queens only take part in a mating flight once in their lifetime, due to
queens being able to store up to 100 million sperm in her oviducts, which she
will use throughout her life to fertilize eggs (Wilhelm
et al., 2010). The queen can also control the sex of her offspring by
choosing whether to fertilise the eggs or not, if fertilised the egg will
become a female worker which are fed royal jelly in the first two days and then
worker jelly the following days, the females are also able to lay infertile
eggs which will become drones. If the egg stays unfertilised it will become a
drone being fed solely on worker jelly (Pechacek et al.,
2005:Peso et al., 2012).
Ant social system
Ants (Formicidae), like termites and bees are eusocial
animals; they also have different castes which have diverse job roles within
the colony (Sempo, 2006).
Workers: Depending on the age of the
workers they will have different jobs roles; the young workers will stay deep
within the nest and the older they get the closer outside the nest they will
get, with older ants working outside the anthill. Workers are female and
sterile, their jobs include looking after the queen, brood, foraging for food
as well as maintaining and expanding the nest (Chittka, Wurm, & Chittka,
2012:Keller, Peeters, & Beldade, 2014).
Soldier: these are sterile female ants
which are larger and stronger than worker ants. Their main function is to
protect the colony from predators and to defend the colony if under attack;
they also use their large mandibles to cut and carry larger objects then the
worker ants (Chittka, Wurm, & Chittka, 2012:Keller, Peeters, & Beldade,
2014).
Drones: the males within the colony,
their primary function is to mate when it’s time for reproduction the drones
will take part in nuptial flight then die as their function has been completed
(Chittka, Wurm, & Chittka, 2012:Keller, Peeters, & Beldade, 2014).
Queen: the queen is the founder of all
ant colonies, she begins her life as a princess (unfertilized virgin queen)
once she has mated, she will stay fertilized for years laying thousands of eggs
daily (Chittka, Wurm, & Chittka, 2012:Keller, Peeters, & Beldade, 2014).
Ant mating system
Ants mating system differs in species,
queens use polyandry. The majority of females can reproduce asexually by
producing unfertilised eggs (Pearcy, 2004). Some male species can plug the female’s
genitals after mating such as the fungus-growing
ants (tribe Attini) (Baer, 2004). In the majority of species only the queen and
breeding females are able to mate. Some colonies have only one queens some had
multiple mates and some survive without any queens (Pearcy, 2004). In colonies
without queens, workers with the ability to reproduce are called gamergates.
Ants with gamergate colonies practise monogyny (D. rugosu) or polygynous (O. hottentota) (Ware et al,
1990).
During nuptial nights, females and winged males
will depart the colony. Nuptial flights happen between spring and summer as the
hot weather makes it easier for the ants to fly, alongside the humidity making
it easier for the queens to dig nests (Dalecky, Debout, Estoup, McKey,
& Kjellberg, 2007).
Males take flight before females, then release a mating pheromone which
attracts the females. Some species of females mate with males only once, while
other species of females can mate with ten or more different males within in
the nuptial flight and store their sperm in the spermathecae for future use
(Zinck et al., 2007).
Mated females will then begin their search for a suitable place to start their
own colony, they then begin to lay and care for their eggs. Females can choose
to fertilise eggs in the future with the sperm they have stored, or lay
unfertilized eggs which will form into workers ants (Keller & Passera, 1992:Timmermans,
Grumiau, Hefetz, & Aron, 2009).
Model one
Kin selection theory provides answers
to the reason behind natural selection favouring cooperation within social
insect societies. Hamilton (1964) argues that kin selection is very important
within insect colonies. Kin selection is a gene which produces copies of itself
either by increasing the fitness of its bearer (direct fitness) or increasing
the fitness of relatives which share the same gene (indirect fitness) which
form the basis of inclusive fitness. The theory behind kin selection is that
helping relatives is beneficial, whereas harming them is not. Alongside when
should an individual help a relative? Hamilton’s rule predicts when altruistic
acts will be favoured by selection (Eberhard, 1975). Hamilton’s rule = r B >
C, suggests that altruistic behaviour will be favoured when the fitness gained
for the recipient (B) compared to the relatedness (proportion of shared genes)
(R) outweighs the fitness loss to the actor (C) (Michod
& Hamilton, 1980:Grafen, 1980).
Kin selection is common in social insects. Several studies have proven that
social insect colonies are family groups (Crozier, 2001). In ants altruism is
contained because there is minimum aggression within their networks due to
worker ants being too specialized to change for a reproductive role. This can
explain the maintenance of altruism within colonies; sterile workers have zero
relatedness and therefore lose heritability. Sterility can evolve due to kin
selection, as sterility gene is only expressed in females which are poorly fed this
causes them to assist well-fed relatives (Eberhard, 1975:Hamilton & Michod,
1980).
|
Regarding Hamilton’s life-for-life
coefficients suggesting that twice as many genes get transmitted by females
compared to males (Hamilton, 1972). Figure
one shows the relatedness the female (Self) shares with her family members. Concerning
the individual (self) the female has 50% chance of sharing the maternally
allele with her full sister, but due to her father only having one allele this
is always shared with her full sister (Richards,
Packer & Seger, 1995).
Kin selection states that the
frequency of altruism increases the population as a result of the behaviour, although
the reproduction of the altruist decreases (Dawkins, 1989:Zahavi, 1995). The
misconception of this model is suggested by Haldane (1995) who proposes that if
two brothers are walking beside a river and one falls in and is in danger of
drowning. It would be advised that the other brother takes the risk of saving
him (decreasing his fitness) in order to increase the frequency of his genes
getting passed on to the following generations.
The model becomes inconstant when told with three or more brothers, as if one brother falls into the river the
other brother who does not risk himself gains as much as the brother which
risks himself but doesn’t occur any cost. The problem with multiple rescuers
shows an unrealistic model as there is no interaction amongst the brothers as
who will rescue the drowning brother (Eshel & Motro, 1988). This suggests that the model only shows
benefits when used within two individuals as the costs and benefits are easy to
predict between two individuals, when more than two individuals are used the
model becomes unclear (Smith, 1964).
Model two
|
The arrows show links and feedback loops, representing the effects due to the influences of:
1.
“colony
size on queen replacement and worker policing
2.
policing on
colony productivity
3.
nonreproductive worker specialization on morphological skew
4.
morphological skew on divergence of queen and worker
lifespans
5.
morphological skew on queen and worker productivity
6.
colony size on optimal system of division of labour
7.
worker polymorphism on colony efficiency and hence colony
size
8.
worker reproductive potential on permissible worker
morphologies
9.
morphological skew on the timing of caste determination and
hence degree of worker polymorphism.” Quotation from Bourke (1999).
The limits of colony
size are likely to be due to ecological factors, but the change in colony size
creates drastic social consequences. This helps to understand why small
societies have simple organisations, and large societies are often more complex
in their dimorphism between workers and reproductives (Morphological skew) (Bańbura, 1992).
Bańbura
(1992) discusses the connection between colony size, worker
reproductive potential and morphological skews. In small colonies, workers have
a higher chance of replacing the queen because there are fewer competitors and
therefore selection for a new reproductive is higher. This promotes a lack of
specialization for worker behaviour which as a result shows low morphological
skew. However on the other hand, any individual in a large colony has a low
chance of replacing the queen, due to the number of competitors, leading to low
potential of reproducing and increased specialization within worker roles and
resulting in a high morphological skew (Nowak, Tarnita, & Wilson, 2010:Beckers,
Goss, Deneubourg, & Pasteels, 1989). For
example, worker ants have become so specialised within their role that they
have lost their wings and generally all of their reproductive apparatus, and
therefore cannot replace the queen or disperse leading to them being ‘trapped’
into social life. (Schulman, 1978) states that positive links between
reproductive skews and colony size, are not well recognised in existing formal
indices of skews (Sherman, Lacey, Reeve, &
Keller, 1995). Stillwell et al (2010)
argues that increased colony size improves future reproduction regardless of
the higher chance of dimorphism within the cast members.
In regards to the
model, there is not a lot to fault due to the basis being correct, large
colonies do have more complex social structures and smaller colonies have
simpler social structures (Goodisman,
2001). Termites have extremely large colonies with up to one
million individuals in some species; they are very diverse in their castes (Noirot,
1989). Their reproductive caste consists of
primary, secondary and tertiary reproductives showing complex diversity compared
to bumblebees, which nests can contain on average 400 bees. and only have castes of Queen, worker and drones
(Pereboom, Jordan, Sumner, Hammond and Bourke, 2005).
In conclusion, ants, bees and termites
are all eusocial insects ( Varley & Klopfer, 1963), due to being eusocial they all live in large colonies
with a caste system. The castes
differentiate within the three species, however all three species have a queen
and workers within their castes, bees have an additional caste of drones, alongside
termites and ants having additional castes of soldiers. All species being mating during a nuptial
night when the weather conditions are correct they will disperse from the
colony, mate, and find a new nesting site. Termites become lifetime monogamous
once mated, whereas bees and ants take part in polyandry. All three species are
able to reproduce asexually without having to fertilise their eggs. Bańbura
(1992) model
suggests that the larger the colony size the less chance there is for workers
to reproduce, which is why some species have lost their reproductive organs,
the model is correct in the fact that species with larger colonies have more
complex structures. Kin selection also plays an important role in eusocial
animals, suggesting that altruism shown in an individual can increase the
population as a whole. However the model shows flaws when used in more than two
individuals. Overall species social and mating systems can be influenced by a variety
of factors, and thus more research needs to be conducted in order to understand
these species further.
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