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ANIMAL BEHAVIOUR
Mechanisms
Behaviors involved in interactions among species are agonistic behaviors, predator-prey behaviors
and reproductive behaviors.
These behaviors result in individual distance the "kick-kiss" distance.
Spacing patterns and social groupings
Spacing pattern
The order Hymenoptera is the largest and most well-known animal group with eusocial species. Most
hymenoptera are not eusocial, but this social system has arisen multiple times within the group. It is
found in some bees (Apidae) and wasps — with separate origins of eusociality in Sphecidae and
Vespidae — and all ants . Hymenoptera have a haplodiploid sex determination system (whereby
females arise from fertilized diploid eggs and males arise from unfertilized haploid eggs), which may
contribute to kin selection, favoring altruistic behavior in this group.
Termites can be considered to be highly evolved social cockroaches which live in their food. They
depend on a complex mutualism with cellulose digesting protozoans and bacteria, and young
individuals acquire these symbionts via anal trophallaxis. Within a colony, there is caste
differentiation: there is a queen and king, the sole reproducing individuals, and there are soldiers and
workers, which are different stages of pluripotent larvae of both sexes. Termites are diploid and
perform complex social behaviors including nest construction and territorial defense .
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The more recently discovered eusocial organisms include a few species of shrimp, aphids, and thrips.
There are at least two separate origins of eusociality within the Synalpheus shrimps . These marine
shrimp live in groups of several hundred closely related diploid individuals as internal parasites on
tropical sponges. The host sponges have a heterogeneous distribution, and this may have contributed
to the evolution of eusociality within this group as dispersal to establish new colonies is riskier than
remaining within the natal nest.
Thrips are small haplodiploid insects in the order Thysanoptera. Of about 5,000 species, about 300
form nests in plants called galls, feeding on plant tissue within. Of these, there are six species which
can be classified as eusocial: they have morphologically different soldier castes which defend the
galls from kleptoparasites.
Social aphids, like thrips, live in galls or hollowed stems of plants, feeding from the plant tissue
(Stern 1994, Aoki & Imai 2005). These tiny hemipterans may have complex life cycles — they are
diploid, but can reproduce parthenogenetically — and there have been several species described that
have robust soldier morphs .
There are at least two species of vertebrates that could be considered eusocial, the naked mole rat and
the Damaraland mole rat. Both species are diploid, highly inbred and live in harsh deserts with patchy
food resources. Most individuals help to raise siblings or close relatives that are born to a single
reproductive female (the queen). Outbreeding results from the generation of dispersive morphs —
individuals that have inbreeding avoidance and large fat stores — from the natal nest to start a new
colony.
Advantages to Living in Groups
Eusocial organisms live in groups, and are subject to both the benefits and costs of group living.
Living in a group may confer benefits on individuals in several ways. First, groups may form as
defense against predation, forming a "selfish herd" in which each individual has a lower chance of
being killed. Second, larger groups in many species gain advantages against competitors, such as in
colonies of the ant Azteca trigona.
Many eusocial organisms have strategic advantage when acquiring food in groups. Where there are
limited nest sites or resources are patchy, the benefits to staying within the natal group include
reducing dispersal risk and the possibility of inheriting the natal nest. This is a contributing factor in
naked mole rat societies and in insect societies with totipotent workers such as primitive ants and
wasps.
There can be costs to living in social groups, and these must ultimately be balanced by fitness
advantages. Disadvantages include increased competition for resources between individuals,
increased transmittance of parasites and diseases within groups, and easy detection of the group by
predators and parasites.
How did Eusociality Evolve?
Giving up one's reproductive potential is contrary to the basic premise of natural selection (to survive
and reproduce). Even Darwin (1859) commented on the challenge of understanding eusociality as
"one special difficulty, which at first appeared to me insuperable, and actually fatal to the whole
theory. I allude to the neuters or sterile females in insect communities: for these neuters often differ
widely in instinct and in structure from both the males and fertile females, and yet, from being sterile,
they cannot propagate their kind." Darwin goes on to argue that "This difficulty, though appearing
insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be
applied to the family, as well as to the individual. . ." There have been several hypotheses proposed
for the evolution of worker-like behavior. It is important to note that they are not mutually exclusive
— each may play a different role in the evolution of eusociality in different groups. Evolutionary
biologists trace the origins of eusociality through a pathway that starts with solitary organisms
acquiring benefits to group behavior, eventually leading to a "point of no return" wherein certain
individuals no longer have the physical ability to reproduce and only gain evolutionary fitness
indirectly. In addition, it is important to note that the selective forces at work during the inception of
eusocial behavior may be different from those that maintain advanced eusocial colonies. What
follows is a brief description of major contributing factors during the inception of eusocial behavior.
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Figure 4: Diagram illustrating factors contributing to the evolution of primitive eusociality and that
advanced eusociality is a social state that is not reversible (past the point of "no return," individual
workers have lost the capacity to reproduce independently)
Kin Selection
There are two routes by which a gene can promote copies of itself in future generations: directly,
through producing offspring, and indirectly, through the reproduction of close relatives. The sum of
direct and indirect reproductive gains is known as inclusive fitness. As a result, it is possible for
selection of eusociality over solitary behavior if indirect fitness levels exceed direct fitness.
An altruistic act is one which benefits a recipient at a cost to the performer of the act. Hamilton's rule
(Hamilton 1964) states that altruism is favored if
r > C/B
where B is the benefit to recipient of altruistic act in terms of lifetime reproductive success and C is
the cost (decrease in lifetime reproductive success). The coefficient of relatedness, r, ranges from 0 to
1 and is the proportion of alleles shared between two individuals identical by descent. Eusociality
depends on high levels of altruism within groups as individuals increase the fitness of others at a cost
to themselves (e.g., by helping care for brood, defending nests, sharing food resources, not investing
in their own offspring).
There are two well-known mechanisms by which r can be elevated: haplodiploid sex determination
and inbreeding. While is it not required for eusociality to evolve (naked mole rats and termites are
diplodiploid), haplodiploidy may help to explain why eusociality has arisen multiple times within the
Hymenoptera. In haplodiploid organisms, the relatedness between full sibling sisters (r = 0.75) is
greater than between a mother and her offspring (r = 0.5), thus weighting Hamilton's rule in favor of
raising sisters rather than offspring.
Inbreeding, or mating between close relatives, results in offspring which share a larger proportion of
alleles, thus increasing r. This is common in organisms which don't disperse very long distances from
the natal nest or are prone to mating with siblings (e.g., termites and wild naked mole rats;
Delayed Benefits
An intermediate step toward eusociality may have "hopeful reproductives" — that, workers which
have the option to stay and help or to go out start their own nest. The decision can depend on
hierarchical position within the group, territory, or food resources and other environmental
conditions. Florida scrub jay offspring are known to stay at the natal nest, helping to raise siblings
(thus increasing their inclusive fitness) until there is an opportunity to replace their parents.
Primitively eusocial wasp colonies, such as Polistes, are commonly inherited by dominant workers on
the death of a queen.
Multi-level Selection
Natural selection can occur at the level of the individual, family (related individuals, known as kin),
or group (non-related individuals). This is known as multi-level selection. For eusocial organisms, the
traits (phenotype) of the colony interact with the environment to determine colony-level fitness and
can be described with models of multi-level or trait-group selection. Whether models of multi-level
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selection or inclusive fitness models are the best approach to explore the origin and maintenance of
eusociality remains a strong point of debate especially since there is limited empirical evidence for
inclusive fitness in groups .
Ecological and Life History Contributions
Nesting behavior has been described as a possible prerequisite for the development of eusociality, in
large part since it creates situations conducive to cooperative brood care . Where nest founding is
dangerous or there are limited territories or spaces, "fortress defenders" can cooperate to defend this
valuable resource. In the case of termites, thrips, shrimp, and aphids, the protected nest is also the
location of food in a patchy environment. Parental care can also be an important life history
component. One path to eusociality in Hymenoptera is thought to start with solitary females engaging
in simultaneous progressive provisioning — rearing multiple larvae of different ages at the same
time. Transitioning to eusocial behavior would then incorporate remaining offspring and provisioning
siblings, followed by offspring withholding their own reproduction.
Communication Among Animals
Communication is the transfer of information. Animals often communicate among their own species.
Sometimes they even try to communicate with other species. Most communication is done through
body language. Sounds and odors are also means to communicate. Humans can learn from how
animals communicate among each other, since the same methods apply in a subtle manner among
humans.
Questions you may have include:
• How do animals use body language?
• How do animals communicate with sounds and odors?
• What can humans learn from animals ?
• Communication Among Animal Species
• 1. Scent
• Smell is probably the most common basic means of animal communication with even the
most primitive animals reacting to odours given off by their own or other species.
• Animals may use scents to proclaim their readiness to mate, to mark out territorial boundaries,
to warn off intruders and predators or, in some cases, to attract prey. The most basic substance
for these purposes is strong smelling urine, but there are other and more refined methods of
producing lingering odours. These take the form of scent glands which may be situated in the
hooves, on the face or close to the anus or scrotum.
• Communication by scent can prove dangerous to certain animals such as the musk deer and
certain civets. Their scent is so strong and persistent that their glands are highly prized by the
makers of perfumes who use it as a stabiliser.
• Bees, wasps, ants, moths and other insects rely largely on upon pheromones as a means of
communication. Pheromones are chemical substances which may be secreted in urine, dung or
produced by special glands. They are usually given off by the female of the species to attract
males. For instance, the antennae of some male moths are so sensitive that they can detect the
presence of a female at a distance of more than two miles!
• Sharks have a particularly efficient sense of smell and some species can detect drops of blood
in the ocean at a range of about one mile.
• Snakes have the most unusual means of detecting scents. They use their tongue to pick up
scent particles in the air. Having flicked out its tongue the snake then returns it to a special
organ in the roof of the mouth which analyses the scents collected.
Dung is another readily available source of scent used by many animals to mark their
territory, with some species accumulating and regularly adding to large heaps of droppings
which act as territorial marker posts. The rabbit, vicuna, West African civet and hippopotamus
are just four examples of these.
2. Sound
• If the olfactory signs are the most common form of communication among animals the
acoustic signals must surely be the next in line, as animals of all kinds rely to a great extent on
their hearing ability in order to succeed and survive.
• Among mammals, generally speaking, small animals squeak and large ones rumble. The
reason for this is that the smaller the animal's head, the higher the frequency of sound it can
receive and transmit.
• In order to determine which direction a sound is coming from, the brain must distinguish
which ear it reaches first, and the time difference between them. The closer the animal's ears
are together, the shorter the wavelength, and therefore the higher the frequency, will be
necessary for good discrimination.
Kangaroos, hares and rabbits will thump their hind legs on the ground as a warning signal.
The rattlesnake gives a distinctly audible and sinister sounding warning by vibrating its bell-
shaped tail segments while other snakes, lizards and crocodilians will hiss loudly to warn off
intruders.
• The study of acoustic communication among animals is producing new discoveries each year
- leaving us to marvel at the complexity of Nature and the way in which life on Earth has
developed.
• 3. Sight
• Visual signals may take the form of gestures and display, facial grimaces, body posture or
mimicry. The extravagant display of the peacock or lyre bird can make the strutting of a wood
pigeon seem ineffective and yet each species has its own way of using visual communication
to the best advantage.
• The male rabbit will use the white underside of its tail to attract the attention of a female,
while the female may use the white of her tail as a visual signal for her young to follow when
she is leading them to the safety of the burrow.
• Dogs and wolves make use of body language, as do cats, monkeys and many other animals.
The attitude of the tail when two wolves meet will indicate which of the two is the superior.
The tail held between the legs is a submissive gesture while the tail raised confidently aloft
donates dominance. Horses will hold their ears or tail in certain positions to signal pleasure or
alarm. Dogs wag their tails as a sign of pleasure while the waving tail of a cat signals irritation
or anger.
• Apes and monkeys make considerable use of facial grimaces in order to express their feelings.
The frown of a rhesus monkey is a clear sign of unease, whereas the raised eyebrow and
fluttering eyelid denotes friendship or pleasure. Raised hackles in the Dog family help to
make the individual appear bigger when facing a possible adversary.
• At another level animals may communicate a simple message through the art of mimicry. The
roundels on the wings of the peacock butterfly look like large eyes to a potential predator. The
EVOLUTION OF BEHAVIOR
One general view in the study of the evolution of behavior is that behaviors can have a genetic basis.
This is not to say that all behaviors are genetically based; indeed many behaviors are entirely
culturally transmitted or learned and may have little to do with genetics (why are you sitting in the
same seat?). For genetically influenced behaviors we can treat them as we would treat any other
genetically controlled trait of an organism: 1) if there are genetically based differences in a behavior,
and 2) these differences affect fitness then, 3) behaviors can evolve by natural selection.
Two examples of genetically based behaviors: cricket song. Different species of crickets have
different calling songs with different characteristics, e.g., inter chirp interval, pulse repetition rate, etc.
Hybrids between closely related species often exhibit songs with intermediate characteristics
Another example (on a larger phylogenetic scale) is head scratching with the hind leg in amniotes
(reptiles, birds, mammals; those with an amniotic sac). Most reach the hind leg over the fore limb to
scratch the head; that birds and mammals do it suggests that this behavior has a genetically
programmed basis and has been inherited through much of higher vertebrate evolution.
Behavior is usually dissected into two components for analysis: Proximate causes/questions in
which one asks how the behavior is performed and ultimate causes/questions in which one asks why
the behavior is performed. Tinbergen has identified four questions to pose when analyzing a
behavior 1) what is the cause, 2) what is the development (ontogeny), 3) what is the current function
4) what is the phylogenetic history. A strict course on evolution focuses more on the latter two
questions (recall adaptation/preadaptation/exaptation discussion and the identification of current
utility vs. historical origin).
Herring gulls breed is large colonies on the ground and defend territories. Two separate calls used for
1) advertising nest site ("choking" call) and 2) as a territorial claim (the "oblique pose" and "long
call"). The Kittiwake also breeds in colonies but nests on vertical cliffs and its nest pad is its territory
and breeding site. In this species only one behavior serves both functions: "choking" behavior is both
defensive and part of mate recognition/pair formation. This is seen as an adaptive behavioral shift wit
respect to the nest location (steep cliff).
There are many behaviors that at first appearance do not seem "adaptive". Infanticide in lions was
first viewed as "aberrant" behavior by abnormal individuals because it was not "good for the species"
(male lions displace other males from groups of females and their offspring, and frequently kill the
cubs). It is true that killing infants is not, in the short term, an effective means of increasing
population numbers of a species. BUT, we now know (post W.D. Hamilton's 1963, 1964 papers on
inclusive fitness and kin selection and G. C. Williams book on Adaptation and Natural Selection) that
the more appropriate way to address such problems is to think about them in the context of whether
the behavior is good for the individual.
In analyzing infanticide from the perspective of gene thinking it is 1) not adaptive for a male lion to
invest reproductive effort in an individual with whom he shares no genes and 2) once the infant is
killed it is advantageous for the female to come into estrous and have more offspring with the new
male (this will increase her reproductive output over leaving with the displaced male, and not
benefiting from other advantages of group living: foraging, avoiding predation on young). Given the
situation for both male and female, the observed behaviors make sense in terms of propagating ones
genes.
The role of the gene (or genes!) as the unit that is relevant in the evolution play an important part in
two influential books in the mid 1970s Sociobiology by E. O. Wilson, and The Selfish Gene by
Richard Dawkins. To grossly oversimplify one of their main messages: "an organism is just DNAs
way of making more DNA"
If we take the case of bird migration we want to know how the bird navigates to the breeding location
(solar and magnetic cues during flight), how the bird knows when to begin migration (internal clocks
and changes in day length [physiological changes]). There is usually a high cost associated with
migration so we also want to know why birds do it since many die in the process (more time for
feeding, more available food). Individuals that do migrate must leave more offspring than those that
do not - again gene thinking helps account for why the behavior exists
Population genetic approaches to the evolution of traits rarely tell us why a phenotype affects fitness
in a particular way; the models usually look at whether fitness increases or not. The optimality
approach to the analysis of behavior attempts to builds models where different behaviors are treated
as the traits and asks which one of these behaviors might evolve. The approach generally ignores the
mechanics of underlying genetic basis of the behavior (i.e., its mendelian and transmission
genetics). Optimal models assume there is a genetic basis and treat each behavior as a haploid
(asexual) trait that is inherited intact.
While Gould and Lewontin (and many others) have criticized optimal models, the builders of optimal
model (e.g., John Maynard-Smith, Univ. Sussex) argue that the models do not assume that the
organisms are optimal (because there are constraints on evolution of traits), but by treating the
problem as an optimality issue, it can tell you what kinds of behaviors might evolve.
Two general type of optimal models: frequency independent models are designed independent of
what other strategies are doing, and seek to define the conditions which might influence behavior