Animal Behaviour

Animal behaviour: the scientific study of how animals interact with each other and with other
living beings and with the environment. It is a branch of biology that deals with the behaviour of
animals, encompassing fields such as ethology, comparative psychology, behavioural ecology &
sociobiology

Comparative psychology: scientific study of behaviours and mental processes of non-human

animals . Relating to phylogenetic history, adaptive significance & development of behaviour

Animal Science : studying the biology of animals that are under the control of humankind .
Animal cultivation for livestock species

Ethology: The science of animal behaviour. Scientific and objective study of animal behaviour
under natural conditions and viewing behaviour as an evolutionary adaptive trait

Some important people Pioneering Animal Behaviour

It is difficult to look at animal behaviour because we tend to look at it at an anthropocentrism

view (man looking at behaviour). It is important to recognized that some behaviours are
genetically pre-programmed and others are nurtured (learned)

1. Karl von Frisch - (behavior of honeybees) Above all, he has elucidated what has rightly
been called 'the language of bees'.
● Orientation of dance : When a bee has found flowers containing nectar, it performs a
special dance when returning to the hive. The dance informs the position of the sun to the
food source (by means of analysing polarized, ultraviolet light from the sky)
● Length of the waggle portion : distance of food source from the hive
● This is genetically pre-progammed in bees (fixed action pattern)
● However, when it comes to differentiating colours - bees can learn via classical
conditioning as stated by Menzel et al., (2009) whereby free flying bees readily learn to
associate a coloured stimulus with a reward. A single exposure to a sucrose reward on a
colourful plate mediates learning & the formation of STM & experiencing a few colour-
reward pairings leads to LTM colour memory

What is a fixed action pattern?

● Sequence of unlearned behaviour

● Nearly unchangeable
● Carried out to completion
● Sign stimulus (releaser) - produces behaviour
 ● Is an innate behaviour within the species
● These behaviors may be critical to increasing an animal’s fitness

1. Tinbergen used a number of experiments to show fixed action pattern behavior in animals
which will eventually give rise to learning
● Digger wasps uses landmarks around their nest to keep track of the location of
their nests. It was observed that the bee would circle its nest in an ever widening
circle before flying away to hunt. This behaviour was a fixed action pattern - it
was performed the same way each time (instinctive behavioural sequence that is
relatively invariant within the species)
● It was observed that when the landmarks around the nest was manipulated, the
bee would be disoriented when it returns
● The fixed action pattern had given rise to learning; (1) Spatial learning - spatial
structures of the environment, (2) cognitive maps: internal representations of
spatial relationships

Further examples of Fixed Action Pattern:

● The graylag goose: always retrieve an egg that has been bumped out of the nest in
the same manner. The goose will carry this sequence to completion even when the
egg slips away during the process (the goose will also perform the same action
with anything round placed near its nest)
● Three- Spined Sticklebacks Fish: During the breeding seasons, the male fish will
develop a red belly and build a nest. He then courts females with a courtship
dance & encourages her to lay eggs which will then be fertilized by the male fish.
The male will aggressively attack any other males who are near the nesting
territory. Tingerben discovered that male sticklebacks would also attack model of
fish even if they are unrealistic looking (but only if the undersides are painted red
like the red belly of breeding males). When a more realistic model is introduced
that does not have a red belly (appearing similar to a female stickleback), the male
will not attack. It was concluded that the red belly is the sign stimulus that
triggers the attack behavior—the FAP.

2. Konrad Lorenz (Nature vs Nurture)

● Emphasizes the role of imprinting: irreversible learning limited to a sensitive
period in an animal’s life; it enhances fitness by enabling rapid learning
● Not all examples of imprinting involve the parent offspring bonding
● An example is seen in baby sea turtles: According to Lohmann & Lohmann
(2003) hatchlings used visual cues (imprinting) to set and maintain their course
to sea despite being hatch at night. Apparently, the turtles are attentive to light
levels near the horizon (crawling towards low bright lights and ignoring bright
lights) = the sea (reflectance of the moon and stars on the shores)
 ● On beaches shared with humans which has caused light pollution mainly due to
coastal development - posed a serious problem for the turtles as the turtles are
misoriented= predation by other animals

The Birds, eggshells and predator avoidance hypothesis is an example of a nature (pre-
programmed response) occurence where the behaviour seem grounded in instincts . First time
breeders usually remove shells experimentally placed in their nests even before they have laid
their first egg

Reasons for eggshells disposable behaviour increases the likelihood that chicks will survive and
go on to reproduce, making the evolutionary value of this behavior to birds very high:-

1. Sharp edges could injure newly hatched chicks

2. Unhatched eggs could be trapped by eggshells
3. Hatched shells could reduce nest hygiene and increase the risk of bacterial infection
4. Hatched shells could reduce nest camouflage, thereby increasing risk of visually oriented
predators

Nikolaas Tinbergen: The 4 Questions

Are complementary categories for explanations of behaviour. It suggests that an integrative

understanding of behaviours must include both proximate (how/ an event which is closest to, or
immediately responsible for causing, some observed result.) & ultimate (functional) analysis
of behaviour as well as understanding both phylogenetic & developmental history and operation
of current mechanism

Ultimate “why does the behaviour change” Proximate “how does the behaviour change”

Adaptive Value : Behaviours will evolve to Mechanism: how the behaviour appeared in
Selective advantage behaviour (highly accordance to physiology factors such as
functional behaviours) physical morphology, molecular factor
(Causation in the physical construct of the
organism and the behaviour presented)
Example: Lighter wings gives rise to flying
behaviour in birds

Phylogeny: Evolutionary selective factors Ontogeny: Changes in the behaviour of

that had influenced the development of that organism over a lifespan that occurs from
behaviour learning (environmental factors)
Harry Harlow’s Experiment: Behaviour Is Nurtured

● Using methods of isolation and maternal deprivation, Harlow showed the impact of
contact comfort on primate development. Infant rhesus monkeys were taken away from
their mothers and raised in a laboratory setting, with some infants placed in separate
cages away from peers. In social isolation, the monkeys showed disturbed behavior,
staring blankly, circling their cages, and engaging in self-mutilation. When the isolated
infants were re-introduced to the group, they were unsure of how to interact — many
stayed separate from the group, and some even died after refusing to eat.

Dawkins: Selfish Genes Theory

● Selfish gene theory: the more two individuals are genetically related, the more sense it is
to behave selflessly with one another (increases survival and propagation of genes)
● According to Dawkins’ theory, the base unit of evolution is the individual gene.
● Since evolutionary fitness is measured in terms of passing on your genes to the next
generation, Dawkins theorized that each gene acted in a way that would allow it the best
opportunity to be replicated and passed on to the next generation.
● Tit For Tat: Showed how cooperative behaviour might emerge in very simple organism
and maintained as under the right conditions it is an evolutionary stable solution
● When organisms have long life spans, the most evolutionary stable solution is to live in a
social grouping so resources could be shared (increases chances of survival to pass on
genes). Stable payoff equilibrium have been shown for behaviours such as cooperation &
altruism
● Example in altruism: Even Though it does not always benefit the individual, it benefits
the genes and allow the transmission of genes. The most extreme form of altruism is in
eusocial societies (ants) where individual ants give up their freedom to be workers in
order for the queen to reproduce and propagate the genes.

Sapolsky

● Humans and primates are more prone to stress related diseases due to psychosocial
stressors
● Stepwise descent in socioeconomic status (SES) for humans predicts increased risks of
cardiovascular, respiratory, rheumatoid, and psychiatric diseases; low birth weight; infant
mortality; and mortality . (the influence of SES on health)
● In animals (social species), dominance rank influences the extent to which an individual
sustains physical and psychosocial stressors. Thus, dominance rank can potentially
influence an individual animals vulnerability to stress- related disease
● High ranking & Low ranking individuals will experience stress depending on the context
for example:
 1. Resource inequity. In ‘‘despotic’’ hierarchies in which resource access is
skewed markedly and dominant positions are attained through
aggression and intimidation., social subordination in despotic species
can be associated with the greatest physiological indices of stress.
2. Maintenance of dominance. In some species, rank is lifelong and inherited
(for example, in female rhesus monkeys); in others, it may fluctuate. In
species where ranks shift, how does an individual, once attaining a high
rank, maintain it? At one extreme among species with despotic
hierarchies, high-ranking individuals frequently and aggressively reassert
their domination over the subordinate cohort (even in the absence of an
overt challenge). In such species, which include dwarf mongooses,
African wild dogs, and ring-tailed lemurs, dominant individuals have the
greatest physiological indices of stress, most plausibly reflecting the
physical demands of frequent fighting . In contrast, in other despotic
species, high-ranking individuals maintain dominance through
psychological intimidation rather than aggression (where, for example,
mere eye contact with the alpha individual might elicit subordination
gestures). In such cases (e.g., savanna baboons, rhesus and squirrel
monkeys, mice, rats, and white-throated sparrows), subordination is
associated with the greatest physiological indices, plausibly reflecting the
frequent psychological stressors for subordinates and the paucity of
physical stressors for dominant individuals.
● A small literature demonstrates that animals who are more socially stressed by the
dominance hierarchy demonstrate (i) basal hypertension; (ii) a sluggish activation of the
cardiovascular stress response after a challenge and delayed recovery when it abates; (iii)
a pathogenic cholesterol profile; and (iv) increased vulnerability to the atherogenic
effects of a high fat diet
● Chronic stress cause problems in reproductive functions in both males and females. In
females, delayed puberty, decreased levels of estrogen and progesterone, greater risk of
miscarriage & etc. In males, atrophy at testes & hypothalamic regions responsible for
gonadotropin release
● Affiliative behaviour among subordinates can reduce effects of stress (social grooming,
altruism such as caring for another individual’s child)

The Trimates aka Leakey’s Angels

● A title given to 3 women (Dian Fossey, Jane Goodall & Birute Galdikas) by
anthropologist Louis Leakeey. These women pioneered the study of primates in their
natural environment and had contributed significantly to our understanding of primates
(prior to their study little is known about primates)

Jane Goddall

Tool use: the use of an object to

alter the form position, or
condition of another object,
another organism/ the user itself
when the user holds or carries
 ● Jane Goddall on her field observation, discovered that chimps are capable of tool use ( an
ability thought to be only possessed by man)
● Apart for chimps who use vine to rid themselves of flies & parasites, capuchin monkeys
and elephants also demonstrated tool use for this purpose (branches/leaves)
● Goddall also noted discovered that chimps lives in social group and they hunt together
( chimps uses stick to catch termites = tool use)

Birute Galdikas

● Galdikas was the first to observe that orangutans were fruit eaters, and she has since
detailed more than 400 types of food they consume in the wild. She also documented the
long orangutan birth interval, learning that orangutan mothers can spend more than seven
years caring for their young at Tanjung Puting, the national park where she observed their
behaviour.
● While working, Galdikas noted how rapidly the orangutans were losing their habitat
because of deforestation.
● Crisis today: as the result of poaching & habitat destruction, the orang utan population
are on the edge of extinction (Borneo lost 30% of their forests since 1973 & 80% of the
rainforest in Malaysian Borneo had been logged)
● The oil palm crisis: the cheap production costs and growing demands for palm oil have
placed pressure on palm oil producing countries such as Indonesia to rapidly expand their
oil palm plantation & ramp up their production processes. They do this by clearing vast
area of old growth rainforests to make way for their plantation = illegal
● Over 50,000 orang utans on the islands of Borneo & Sumatra have died because of oil
palm deforestation
● A 2007 assessment by the United Nations Environment Program predicts that orang utans
will be virtually eliminated in the wild within 2 decades if current deforestation trends
continue
● Galdikas set up a reserve with a rehabilitation program, and so far, her team is
responsible for the treatment and release of more than 500 wild-born, ex-captive
orangutans back into the wild. Through her foundation, Orangutan Foundation
International, she’s also raised millions in funds to purchase wild forest and preserve
habitat to release her orangutans into.

Dian Fossey

● Studied social structure of gorillas (social structures, hierarchies), vocalization,

eating behaviours and etc

Defining Culture

● No agreeable consensus on how to define culture particularly in animals

 ● One acceptable broad definition of culture by Hoppitt & Laland (2013) - group-
typical behaviour patterns that are shared between members of a community that
is contingent upon social learning
● First evidence of culture (observed in the monkeys of Koshima Island, Japan):- A
female monkey named Imo washing sand off sweet potato & wheat distributed
across the beach by experimenter using sea water. Subsequently, behaviour was
observed in other monkeys
● Transmission of behaviour is thought to happen two ways; (1) vertical
transmission - mother to child, (2) horizontal transmission : from an individual to
peers
● The hundredth monkey effect: hypothetical phenomenon in which a new
behaviour/ idea is said to spread rapidly by unexplained means from one group to
another group

Elephants

Among the Asian elephants, the grandmothers have a significant role. They ensure survival of
the calves & breeding success for their daughters. Research found that calves of young elephant
mothers under 20 years of age had 8 times lower mortality risk if the grandmother resided in the
same location compared to calves where grandmothers were not present.

The team discovered that the more offspring a grandmother elephant has, the greater the chance
of her grand calves' survival. Not only does the grandmothers' presence increase calf survival,
but they also appear to boost the reproductive rate of their daughters in the group.

Musth or must is a periodic condition in bull elephants characterized by highly aggressive

behavior and accompanied by a large rise in reproductive hormones. Testosterone levels in an
elephant in musth can be on average 60 times greater than in the same elephant at other times.

The value of long-term research and individual identification

1) Survival rate - life history strategy

● R strategy: Cheap offspring & unstable environment, lower parental care
● K strategy: Expensive offspring, produce few, live in stable environments, higher
parental care
2) Age group for reproduction
3) Breeding intervals
4) Emigration / mortality rate
5) Population dynamics & Population viability
● Population viability is a species-specific method of risk assessment frequently
used in conservative biology. Defined as the process that determines the
probability that a population will go extinct within a given number of years
6) Society structures
Purpose of Play

In humans, play serves many valuable purposes. It is a means by which children develop their
physical, intellectual, emotional, social & moral capacities, & a means to creating & preserving
friendship

In animals, play is a motor activity performed postnatally that may appear to be purposeless
depending on the inventiveness of the observer;-

1) If an activity is directed towards another living individual = social play (tumble &
chasing)
2) If it is directed towards an inanimate object = object play (manipulation,pulling, pushing
& chewing)
3) If the activity carries the individual in a seemingly frantic flight about its environment =
locomotor play (running, leaping, sliding)

Behaviour & Reproduction

1. Sexual Selection is dependent on:

● Female’s choice
● Male-male competition
2. Sexual selection leads to traits that improves an organism’s chances of reproducing
(certain conspicuous physical traits such as pronounced coloration, increased size/striking
adornments ) but not necessarily survival
3. Secondary sexual characteristics (physical characteristics that distinguishes between the
sexes)

E.g: The Birds Of Paradise: Females choose mates based on condition and colour of the
males’ plumage. Bright & flashy coloring signals that the male is healthy and will
produce healthy offsprings. These traits however make animals more visible to predators.
Therefore, sexual selection does not favour natural selection. In sexual selection,
organisms that have more proficiency in securing mates have higher fitness

In bowerbirds, it is the female’s choice to mate with the male. The male will build a nest
(out of stick) with well decorated display courts such as colourful, natural & artificial
objects which is used for deco = important role in mating. The visitation of female is
influenced by; (1) male size, (2) display rate, (3) no. of deco, Copulation is influenced by;
 (1) male size, (2) painting rate (male masticating plant material and wiping the plant-
saliva mixture onto the inside walls of the bower). Painting rate is positively correlated
with mating sucess (Robson t al., 2005)

Male-male Competition

1. Lek mating - an aggregation of male animals gathered to engage in competitive

display to entice females
● Competition of strength
● Females will observe and choose the winner
● There is a strong association between female preference and male viability which
supports a basic assumption of the good genes model where choosy females
benefit through better vialibility of their offspring

4. Sexual dimorphism: when males and females have different morphology (example size)

5. Some hypothesis/principle for behaviours in mating

Hypothesis/Principle Explanation

The sexy son hypothesis Female’s ideal mate choice among potential mates is one
whose genes will produce male offspring with the best
chance of reproductive success (flamboyant males)

Honest Signal Theory (The A hypothesis proposed to explain how evolution may led to
handicap principle) honest/reliable signaling between animals which have
obvious motive to deceive. The handicap principle suggests
that reliable signals must be costly to signaller. It is
proposed that females prefer males with handicaps (mating
characteristics that reduce survival chances) because
handicaps are indicators of heritable viability (ability to
escape from predators even though it has bright colours)

The bright male hypothesis/ Female choose male in the basis of male characteristics
parasite mediated sexual (example: plumage that shows male is healthy) so she
selection obtains heritable parasite resistance for her offsprings

In birds, carotenoids (red colour) are limited in the

environment so only birds with high quality diet have
bright red plumage

Bateman’s principle The sex that invests the most in reproduction female (e.g:
ovum, pregnancy, nursing = more investment) vs male
(sperm) is a limiting resource that the opposite sex
competes for. Therefore, females can be choosy
 Parental investment - Ronald When sex is outnumbered, more parental investment will
Fischer be put into the outnumbered sex. Therefore equilibrium of
sexes is 1:1

Parental investment theory - The sex making the llargest investment (females) will be
Trivers more discriminating in mating while the sex that
contributes less in mating will compete for access to the
higher investing sex

Strategies of Males in Mating

1. Infanticide: When a new male takes over, the offspring of the previous alpha male will be
killed. Therefore, it increases the likelihood to father more offspring
● Some solution to counter this by females;
● In baboons, (1)pseudo-estrus - faking female signals that they are sexually receptive so
the male believes the child is his, (2) mating with multiple males, (3) social bonding with
protective males
2. Sperm competition: Seen in insects (diverse shapes for male and female reproductive
organs. Key & lock hypothesis - to prevent hybrids). Males have penis shaped like brush
to remove sperms from competitors, Copulation plug - to block sperms from other
competitors
● Females counters this;- (Cryptic female choice where females use
physical,chemical mechanisms to control for a male’s success of fertilizing the
ovum)
3. Sneaky male: A strategy used by aquatic organism (cuttlefish)- Portrays sexual mimicry
whereby small males will look and behave like females to gain access to females and
avoid aggression with the alpha/dominant male (Evolutionary Stable Strategy)

Behaviour & Society

The ecological niche of a species therefore includes not just the species habitat but also ways in
which it interacts with other species and the physical environment

Predation

The relationship between the Canada lynx (Lynx canadensis) and the snowshoe hare (Lepus
americanus) is considered a classic example of how interactions between a predator and its prey
can influence population dynamics of the two species. Canada lynx populations rise and fall with
fluctuations in populations of snowshoe hares. Thus, when hares are abundant, lynx populations
expand, and when the density of hares is reduced, lynx are forced to hunt ground squirrels,
grouse, and foxes. Though they occasionally kill larger animals such as a white-tailed deer, the
shift away from hares takes its toll, and lynx populations ultimately shrink.This can lead to
cyclical patterns of predator and prey abundance, where prey increase in number and then, with
abundant food, predator number increases until the predators begin to suppress prey numbers and
then decrease as well. As long as predator and prey numbers don't drop to zero, this cycle can
repeat indefinitely.

Anti-predator Adaptation

1. Avoiding detection
● Staying out of sight
● Camouflage
2. Warding off attack
● Startling the predator
 ● Pursuit-deterrent signals- Communication Between Prey. and Predator. When
encountering predators, prey animals often expose themselves by loud
vocalization, by repeated movements or by revealing conspicuous colors
● Playing dead
● Distraction
● Mimicry and aposematism- Conspicuous colors are often used by prey to
advertise their toxicity to predators. Some species have evolved similar color
patterns to toxic prey, capitalizing on the tendency for predators to learn to avoid
conspicuously signaling toxic prey.
● Defensive structures
3. Safety in numbers
● Dilution effect - chance of an individual being captured by a predator decreases
as group size increases
● Selfish herd - individuals within a population attempt to reduce their predation
risk by putting other conspecifics between themselves and predators
● Predator satiation - prey briefly occur at high population densities, reducing the
probability of an individual organism being eaten.
● Alarm calls
● Improved vigilance
● Predator confusion
4. Fighting back
● Chemical defenses
● Communal defenses
● Defensive regurgitation
● Suicidal altruism
5. Escaping
● Flight
● Autonomy - casting off a part of the body

Grouping

● Living and staying in groups is an evolutionary stable solution (see Richard Dawkins)
● The game theory: Interactions between organism range from antagonistic to cooperative.
Interactions are costly depending if no resources is obtained. Organism with the best
interaction strategy (maximizing net gain over net loss) has the highest fitness. Since
biological interaction involves 2 or more decision markers, biologist utilized game theory
to demonstrate evolutionary consequences of interactions. Game theory investigates the
outcome of multi-individual interactions, in which the payoff for an individual depends
on strategy of other individuals involved. Stable equilibrium payoffs have been shown for
cooperation & altruism

Landscape Of Fear
 ● The “landscape of fear” hypothesis predicts that the expression of antipredator responses
reflect a tradeoff between risk avoidance and energy gain, such that animals avoid areas
of high perceived predation risk even if forage is abundant in those areas = change in
ecosystem
● Example: After wolves were reintroduced to Yellowstone National Park in Wyoming in
1995. Elk that had previously browsed in open areas began to avoid them, and willows on
streambanks and floodplains thrived
● Because top carnivores such as wolfes, tigers, lions and etc are capable of killing and
eating mesopredators (coyotes, red foxes, baboons), these mesopredators often avoid
areas claimable to top carnivores and alter their hunting and foraging patterns to reduce
the chances of an encounter (landscape of fear). However in the absence of large
predation, mesopredators can claim the hunting grounds = mesopredator release. The
increase in mesopredator populations can force sudden changes in the structure of
ecosystems (predation pressure on prey such as song birds & small mammals) causing
many such species to suffer dramatic decline= lower levels of biodiversity.
● One notable example of mesopredator release occurred in North American during the
20th century when cougar and wolf populations declined dramatically in the
conterminous of the United States as a result of overharvesting. With these top carnivores
gone, a major source of mesopredator mortality was removed

Mesopredator release hypothesis:

● Ecological theory used to describe the interrelated population dynamics between apex
predators (a predator at the top of a food chain, with no natural predators) and
mesopredators (a medium-sized, middle trophic level predator, which both preys and is
preyed upon) within an ecosystem such that a collapsing population of the former results
in dramatically increased populations of the latter.

Interspecific Competition

● The competitive exclusion principle states that if 2 species with identical niche (i.e:
ecological roles) compete, then one will inevitably drive the other to extinction
● An example: The local extinction of Common myna in Singapore and the replacement to
Javan Mynas. The Javan Mynas were more adept than the common mynas (came out to
forage for food earlier, and is said to be more opportunistic and aggressive). Therefore
Common Mynas which were common in the past were outcompeted by the Javan
Mynas= Javan Mynas taking over the common niche

Birds: mixed species foraging flocks

All around the world, mixed species flocks of foraging birds are thought to forage
together for either improved feeding efficiency/better protection from predators. Sridhar
et al., 2009 found that mixed species flocking benefits species that are vulnerable to
predation. Therefore, they follow species whose vigilance they can exploit. Theodore et
al., 1985 however found that mixed species flocking in birds is due to increased feeding
efficiency alone or feeding efficiency and predation pressure in combination
 Interspecific altruism

Pittman & Schulman-Janiger analyzed 115 rescue incidents involving adult humpback
whales from published and unpublished sources. Only 11% of the time were the
humpback whales trying to save their own calves, the remaining of the time, humpback
whale will rescue other animals such as harbor seals, gray whales & California Sea Lion.
Below are some observations of humpback whales attempting to rescue another animal

1. Pod of killer whales knocks a seal off an Antarctic ice floor. The humpback whale
proceeded to lift the animal and placed it on its belly, keeping it out of harm’s
way
2. Another observation was in Monterey Bay, California in May 2012, when a pod
of killer whales targeted a gray whale and killed the whale’s calf. When the killer
whale first attacked the gray whales, 2 humpback whales were already at the
scene, but after the calf was killed, 14 more humpback whales arrived at the
scene. They vocalize loudly and slashed their tails every time the killer whales
attempt to come closer. Despite the thick swarms of krill nearby - the humpback’s
main choice for food they did not abandon the dead calf

Intraspecific Competition

Happens when:

1) Competing for food

2) Spread of parasites and diseases
3) Density dependent (K strategy)

R Strategy K Strategy

Reproduce rapidly Reproduce slowly

Short lifespan Longer lifespan

Large number of offspring Small number of offspring

Smaller bodies Larger bodies

Lived in unstable environment Found in stable, established habitat

 Strategies To Avoiding Conflict

1) Territory partitioning
2) Dominance - Alpha male & Alpha female
3) Allomothering
4) Cooperative Breeding
● Resource partitioning: Often used for explaining how different species live together in the
same habitat (because they utilise different niches = contribute to the environment
differently). Therefore spatial partitioning & temporal partitioning
● Males & females could live together as their different behaviour cause them to
accommodate different niches.

Predator-Prey Interactions & Anti-Predator Behaviour

● Anti-predator behaviour - suite of behaviours that animals display in response to

predatory threats.
● Behaviour varies with context and does not need to be species specific. Examples of anti-
predator response; counter-attack, flight, refuge use, alarm calls
● Cheney & Seyfarth (1980): Vervet monkeys issue different types of alarm calls for
different predators (Vocally distinct sounds to indicate different predator such as eagle,
leopard and etc). When alarm calls that are recorded was played back to the monkeys at a
different time, different responses were elicited in accordance to the predator. Example:
ran into trees for “leopards”, looked up for “eagles”looked “down” for snake. Infants
particularly gave many calls to mammals, birds and snakes
● Passive observations of behaviour: made possible with technological advances such as
passive infrared triggered video camera, “trailcams
● Thuppil & Coss (2013): played back tiger and leopard growls to Asian elephants in
southern India along paths that elephants used to raid crops in village. Aim: (1) to
investigate if the elephants made different anti-predator response to tiger and leopard
growls because tiger is seen as a threat but not leopards (tigers can opportunistically prey
on elephant calves), (2) determine if the playblacks of these predator vocalizations can be
reliably used to reduce human-wildlife conflict (prevent elephants from raiding crops &
villages)

Results : Elephants produces different anti-predator responses to both leopards and tigers. For
leopards, elephants typically react aggressively (trumpets & grunts). For tigers, elephants retreat
quickly. In conclusion, elephants are able to discriminate between vocalizations to different
species of animals and assess them to be posing different levels of threats (elephants considered
tiger growls to be more threatening while leopard growls were considered less threatening)

Coevolution
 1) With predator-prey relationships, both predator and prey will influence each other
(evolutionary-wise) ;-
■ i)Predators adapt to prey
■ To acquire food, they have to overcome prey’s anti-predator
response
■ ii) Prey adapt to predators
■ Anti-predator behaviour involves dealing with predators
adaptations for exploiting them as prey
2) An example of a co-evolution case study: California Ground Squirrels &
Rattlesnakes developed evolutionary arms race whereby an adaptation in one of
these species will trigger the evolution of an adaptation in the other;- The
rattlesnake's adaptations include toxic venom as well as a specialized infrared-
sensitive pit organ on its face that can sense heat, and thereby help the animal find
prey.To counter rattlesnakes, California ground squirrels developed a rare ability
to neutralize rattlesnake venom.

Adaptation Of Squirrels Adaptation Of Snakes

Venom resistance (snake targets baby Infrared sensitive pits to detect hits from prey
squirrels) (can detect prey in dark burrow)= important in
predatory behaviour

Knowledge about snake habits ( Squirrels Rundus et al., 2017: California ground
know that snakes retreat or hide in burrows = squirrels add an infrared component to their
more cautious around burrows) snake-directed tail-flagging signals when
confronting infrared-sensitive rattlesnakes ,
but tail flag without augmenting infrared
emission when confronting infrared-
insensitive gopher snakes . Experimental
playbacks with a biorobotic squirrel model
reveal this signal's communicative function.
When the infrared component was added to
the tail flagging display of the robotic models,
rattlesnakes exhibited a greater shift from
predatory to defensive behavior than during
control trials in which tail flagging included
no infrared component.

Instigating snakes to rattle by confronting Snake adaptations behaviorally to squirrels

with sand kicking, attacking & killing, •Lying in burrows is an easy way to get a
confrontation elicits rattling = squirrels squirrel to run right into your mouth
discovered cues in this sound for •If squirrels are distracted by other matters,
dangerousness feeding and bobcats and hawks, they may do
just this
• Snakes know when it runs into a female that
really stands its ground =Then it must have
babies nearby
•Non-mothers and males do not stand ground
•Snakes will then look nearby this female
•Use tail-flagging from mom as an indicator
of whether or not they are getting close to
squirrel’s burrow

Using rattle cues, squirrels judge snake size

(bigger=more dangerous they strikes are
faster, longer, and they hold on longer),
Detecting temperature , Temperature= snakes
are ectothermic, cold snakes less dangerous
strike slower, less accurate, less ready to
strike

Physiological effects:
•Squirrels are immune to venom in most
cases. But in 1 area, the venom has changed to
become more damaging to the squirrel;-
Clotting
•Generally, rattlesnake venom causes prey to
hemorrhage to death
•Breaks down blood vessels, tissue, and
ruptures red blood cells
•Components of blood and tissue:
i)Collagen - protein
ii)Gelatin - protein
iii) Red blood cells
iv)Fibrin gives strength to blood clots (More
fibrin = more clotting = less likely of
hemorrhage to death)

= Snake venom takes advantage of the inbuilt

mechanisms (squirrel’s serum) within the
squirrel to break down fibrin
Evolutionary Arms Race In Humans

● Humans did not develop the agricultural settlements until we migrated out of sub-Saharan
African
● Agricultural settlement probably did not evolve then because of animals resistant to
domestication due to an evolutionary arms race that encourages wariness towards bipedal
animals including humans
● Agricultural settlement = domestication of animals that is needed for agriculture purpose
such as cows, buffalos, horse and goats (asian species)
● Research by Brubaker & Coss (2015);- horses will habituate to a great extent to an
approaching humans . However zebras although will habituate as well but only up to a
certain point (distance about 35 m which incidentally coincides with the range of thrown
spears by our ancestors)

In plants (considered predation)

● A lot of similarities because plants defend themselves;- toxins
● Originally evolved as a plant’s defensive compound
- In Woolly spider monkeys, who consumed plants compensates this by eating a few
leaves from different plants = detoxify plants defensive toxins
- In plague bees (overwhelm plant’s toxic defenses). Costly for plants to produce toxins =
only releases it when it is attacked. One way to counter this - lots of beetles attack at
once. In the time it takes the plant to release its toxin, each beetle gets to eat some and
each individual beetle only gets a little toxin
- Tool use to deal with mechanical plant defenses such as nuts with thick shells - chimps
overcome this by using tools, mangabey monkeys do not= culturally evolved mechanism

Predation of animals

• More complicated as animals can behave (they can perceive your attack and can counter your
efforts). Predators can overcome this by;-

Getting around Access prey’s Group hunting Capitalize on Prey’s

prey’s behavioral protected location Perceptual System
defenses

Example: shorebirds Prey defend Example: Harris Example: Orchid

and invertebrate themselves by hawks- Group mantis -
prey (Invertebrates choosing locations hunting in dessert Camouflaging oneself
live in mud under that is hard for so goes undetected by
water, when tide goes predators to spot esp Chimpanzees prey
down, the birds go during vulnerable •Role differentiation
out = pick them out times such as in group hunting Death adder- mimic
of the mud). breeding •In contrast with the prey’s own prey =
However, dangerous: harris hawks attract the prey, attack
 trade off between •Not everyone in it when its close
getting food and own animal behavior enough
safety. Shorebirds are community is
usually in flocks = convinced of this role
lots of eyes to watch differentiation
for predators •But there is some
(falcons). Also lots of evidence to support it
potential prey for •West African chimps
predators (selfish- are better at catching
herd theory) these adult monkeys
than the East African
chimps
•Group hunting
probably has
something to do with
that

Redshanks (hunting Example: sea lion

alone), and is food to nursery beacher.
invertebrate. If in a Killer whales
group, invertebrate (predator) will beach
prey would be scared themselves to gain
off by the redshanks’ access to prey
feet = would burrow
too far into mud.
Venturing out in the
open sand alone to
get food exposes
them to predators

Higher Cognitive Intelligence

1. Cognition: mental processes such as perceiving, recognising, conceiving and reasoning. It

is the mechanism of how you learn, remember and pay attention
2. Intelligence: A high capacity for learning and understanding. An Intelligence Quotient-
test one’s general ability to solve and understand concept
3. It is believed that there is a correlation between encephalization quotient and intelligence

Woflgang Kohler

● Studied the behaviour of apes - developed one of the first cognitive theory of learning
called insight learning ( a type of learning/ problem solving that happens all of a sudden
through understanding the relationships/ various parts of a problem rather than through
trial and error
 ● Experiment: Kohler hung a piece of fruit just out of reach of each chimp. He then
provided the chimps with either 2 sticks/ 3 boxes. Kohler noticed that after the chimps
realized they could not simply reach/jump up to retrieve the fruit, they stopped. In the
first scenario, the problem was solved by placing the smaller sticks into the longer stick
to create a very long stick that can be used to knock down the hanging fruit. In the 2nd
scenario, the chimp would solve the problem by stacking the boxes on top of each other
which allowed them to climb up to the top of the stac of boxes and reach the fruit

Interspecies Communication

● Project Koko was pioneered by Dr Penny Patterson and uses interspecies communication
- establishing a shared language as a tool to understand the inner workings of the great
ape’s mind. A shared language enables us to develop closer, more trusting relationships
with them which in turn leads to deeper (mutual) understanding. While many forms of
interspecies communication is possible with the great ape, sign language has been the
primary vehicle used in Project Koko. Learning based on: Vocabulary - Differentiation -
Transfer - Communication
● Koko’s Gorilla’s Sign Language learning was most rapid (in terms of vocabulary gains)
occurred between years 2.5 & 4.5 years as compared to human children who spike
between 2 & 4 years. Most of the children acquired their first 50 different sign within a
period of 10 months (using emitted criteria). Meanwhile, Koko took 13 months to attain a
50 sign vocabulary. Koko managed to learn over 200 new sign vocabulary in her 3rd year
& her rate of learning tapered off (like humans). Although she continued to acquire new
signs each year, it is not as rapid as human children. Koko’s current vocabulary exceeds
1100 signs. Meanwhile, Michael (Koko’s younger male companion), learned over 600
signs at a similar pace
● Evidence that this ability is not due to the “Clever Hans” effect or any sort of
conditioning;- The gorillas are able to invent a sign for a word by compounding 2/more
sign of pre-existing vocabulary. Example: scratch & comb = brush

Object Permanence

● Defined as the understanding that an object exists even when it is out of the sight. Can
only occur if a mental representation of the object (schema) is formed
● The mirror test/mask test/ mirror self-recognition test developed by Gallup (1970) is a
method for determining whether a non-human animal has the ability for self-recognition
● Found that animals such as the Asian elephants, the great apes, bottlenose dolphins have
passed the mirror test
● De Waal et al., (2006): MSR thought to correlate with higher forms of empathy and
altruistic behaviour . Typical response for an animal that possesses MSR:
(i)social response, (ii)physical inspection (looking behind the mirror), (iii)repetitive
mirror-testing behaviour
 ● The classic mirror test of self-face recognition starts with a mark placed on a subject’s
body. To pass this test, they would have to identify the mirror image as themselves, then
they would have to inspect their bodies to find the mark. Finally, they would have to
realize that the mark shouldn’t be there and attempt to remove it
● Problems with the mark test: difficult to apply across all species for example, primate
would be interested in such thing (groomers) but elephants are different (used to rolling
in mud and etc). Secondly, gorillas generally avoid eye contact as eye contact is a thorny
issue for gorillas and often lead to fight therefore most don’t pass yet they are more
phylogenetically related to humans compared to chimps. Third, not all chimps are known
to pass the mark test - Povinellii et al., (1993) reported that only 21/105 chimps passed
the test= calls into sensitivity of this measure for self-face recognition
● Alternative to the mirror testing: New body awareness paradigm for testing an animal’s
understanding of its place in the environment. The paradigm requires the Asian Elephant
to step on a mat and pass a stick attached by a rope to the mat to the presenter. In order to
succeed in this task, elephants had to see their body as an obstacle and first remove their
weight from the mat before attempting to transfer the stick. Results: all elephants succeed
at least once in the 1st session of the test condition with 8/12 elephants successfully
getting off the mat & exchanging the stick mat at least 11 of 12 test trials

Theory Of Mind

● Understanding that other people’s thoughts and feeling may be different from your own.
Ability to recognise and attribute mental states (thoughts, perceptions, desires, intentions,
feelings) to oneself and to others and to understand how these mental states might affect
behaviour

● 3 form of consciousness;-

Primary consciousness Higher-order consciousness Affective consciousness

Core consciousness/ feeling Extended Core emotional feelings in

consciousness consciousness/self animals & humans
awareness

Moment to moment Awareness of one’s self as A comparatively intrinsic

awareness of sensory an entity that exists function of the brain,
experiences separately from other shared homologously by all
entities mammalian species

Emotions Seeking fear, lust, rage,

care/ nurturance, panic
(separation distress) &
play/social affection

● Cambridge Declaration: Absence of a neocortex does not appear to preclude an organism

from experiencing affective states. Non human animals have the neuroanatomical,
 neurochemical & neurophysiological substrates of conscious states along with the
capacity to exhibit intentional behaviours = consciousness

Ability For Empathy

● Plotnik et al., (2014): Elephants affiliated significantly more with other individuals
through directed physical contact & vocal communication following a distress event.
Suggests emotional contagion (spreading of emotions). Similar consolation responses by
apes

● Spindle neurons/ Von Economo neurons: found in humans, great apes and cetaceans
(humpback,fin, killer & sperm whales). Located in the anterior cingulate cortex,
frontoinsular cortex, dorsolateral prefrontal cortex = active in social reasoning (intuition),
empathy, emotions and others which is in turn important for self monitoring & social
awareness

Evolution of morality (i.e: monkeys reject unequal pay)

● Brosnan & de Waal (2014): Passive and active protest against receiving less than a
partner for the same task is widespread in species that cooperate outside kinship and
mating bonds

● Sensitivity to inequality offers several evolutionary benefits. First, animals need to

recognize when they receive less than a partner, because this tells them that the benefit of
cooperation may be in danger. By protesting against this situation, they show a response
known as inequality aversion. Evidence indicates that this behaviour is widespread in
cooperative species under many circumstances. As the reliance of cooperation increases,
individuals also benefit from sensitivity to receiving more than another, which risks
undermining cooperative partnerships. Pressure for increased cooperation combined with
advanced cognitive abilities & emotional control allowed humans to evolve a complete
sense of fairness
A mixed bad: behaviour manipulators

The behavioural manipulation hypothesis posits that parasites can change the behaviour of hosts
to increase reproductive fitness. Some examples of behaviour manipulators;-

1. Parasite: Toxoplasma gondii

● Infection results in rodents becoming unafraid of cats and instead become attracted to
cat’s urine= higher chances of being predated by cat. Cat becomes infected= sexual
reproduction in cats intestines. Grazing animals ingest the oocytes (from cat’s faeces).
Parasite is passed on to humans by meat
● T.gondii’s preferential localisation is within amygdala (alters fear response)
● Subtle behavioural & personality changes may occur in infected humans. The parasite
has recently been associated with a number of neurological disorders, particularly
schizophrenia and bipolar disorder
● Poirotte et al., (2016) - chimps who are lose their innate aversion towards the urine of
leopards (predator)= increasing the probability of the chimps being predated by leopards
for the parasite’s own benefit
2. Zombie fungus (Ophiocordyceps unilateralis)

● Main goal of ophiocordyceps unilateralis fungus: self propagation & dispersal.

● The fungus manipulates the brain of ants to move to suitable habitat in order for it to
flourish and spread its spores
Timeline:
1. Infects foraging ants through spores that attach and penetrates the exoskeleton & slowly
takes over the exoskeleton & slowly takes over behaviour
2. The ant then leaves its nest for more humid climate that is favourable for the fungus’s
 growth(about 10 inch off the ground). Ant then sinks its mandible into leaf vein and die
3. Fungus feed on its victim’s innards. Several days later, the fungus sends a fruiting body
put through the base of the ant’s head - spread more fungus

3. Hairworms (Nematomorpha)
● Larvea infect hosts (mostly crickets). Once matured, it will drive the host to behave
erratically - jump into water (suicide) - so the worm can swim out

4. Slave- making ants

● A queen from a slave making ant species can take over other ant colonies by using
chemicals to manipulate workers of other ant species to care for her brood
● Slave making ants are brood parasites that captures broods of other ant species to increase
the workforce of their colony. After emerging in the slave-maker nest, slave workers
work as if they were in their own colony

Brain macroorganisation-truine model proposed by Paul MacLean 1960s

Layer 1 -Ancient part of the Layer 2 : More recently Layer 3: recently evolved
brain evolved

Mediates automatic, Emotions. E.g: smell layer of neocortex (upper

regulatory functions. E.g: predator= activate layer 1 to surface of the brain)
body temperature, muscle initiate stress response Primates devote more of their
shiver, blood glucose See something terrifying: brain to this layer (cognition,
activate layer 1 to shiver with memory storage, sensory
emotion processing, abstractions and
etc. E.g: Scary part of a horror
The limbic function = central movie activates layer 3
to emotion which signals layer 2 to feel
Amygdala, hippocampus, frightened and prompts layer
septum 1 to initiate shivering

Neuroendocrine axis

E.g: Stress
Hypothalamus (corticotropin-releasing hormone CRH) ↔ Pituitary (adrenocorticotropic
hormone ACTH) ↔ general circulation ↔ peripheral glands (adrenal glands=
glucocorticoids)

E.g: Reproduction
Hypothalamus (gonadotropin-releasing hormone)↔Pituitary (luteinizing hormone)↔general
circulation ↔ peripheral glands (ovaries=estrogen)
Endocrinology
-study of hormone (a chemical messenger released from secretory cells in various glands).,
Once secreted, it enters the bloodstream, where it can influence any cells throughout the
body that possess receptors for it (receptor ↔ hormone)
-hormonal effects emerges over hours to days and can persist forever (e.g: puberty)
-depending on the hormone & target cells- hormone can change the activity of particular
proteins, turn certain genes on/off, alter the metabolism

-Two main hormones:

Dopamine Oxytocin & Vasopressin in paternal males

Released by the dopaminergic system (ventral For female, stimulates uterine contraction
tegmental area to limbic system & frontal during labour & milk production, facilitates
cortex) maternal behaviour & bonding between
couples

Responds to reward (response determine by Makes organism less aggressive, more

reward value). Once reward contingencies are socially attuned, trusting & empathetic. Foster
learnt, dopamine is less about rewards than social interest and competence= better at
about anticipation. It is about expectation & detecting happy faces/words with positive
confidence social connotations (prosocial) . Enhances
activity in the temporoparietal junction
(Theory of the Mind)

Is bidirectional : dopamine decreases Recent studies however have found negative

following outcome devaluation increase effects of oxytocin whereby people who were
responding when the outcome is not devalued given the hormone and then had to play a
game had more envy & gloating. Kemp &
Guestella suggest that instead of supporting
all social emotions, it plays a role in
promoting approach related emotions
(wanting something instead of shrinking
away)

Influences goal oriented task

Non-invasive Monitoring Of Wildlife Using Faecal Endocrinology

● A new non-invasive analysis of fecal steroid metabolites (estrogen,androgen,

progesterone & glucocorticoid metabolites have been developed
 ● Chester Zoo had used this method of advancement in endocrinology to increase the
mating success of rhinos whereby by monitoring progesterone in these samples using an
enzyme called immuno-assay, the female’s fertility period could be determined =
introduction of male
● Can also determine the levels of stress in animals. For example; it is used in monitoring
stress in Asian Elephants in Peninsular Malaysia
● Extraction of glucocorticoid hormone can be used as an indicator of the elephants’s
health (the hormone plays a role in modulatory daily energy needs & in helping to
prepare the body to cope with challenges, managing the periods of stress & in recovering
after challenge has passed)
● Benefit of using non-invasive methods: Faecal can be easily obtain as wild elephants
defecate up to 18 times a day, animal welfare, information management
● One way to combat the difficulty in observing elephants in the rainforest is to attach GPS
satellite & collaring
● Translocation = moving elephants from conflict zones to protected areas
● Wong et al., 2016 :Translocation of wild Asian elephants (Elephas maximus) is used
extensively to mitigate human-elephant conflict (HEC) in Peninsular Malaysia since
1974. Very little is known about the fate of translocated elephants after relocation due to
challenges in observing elephants in the dense rainforest. Advances in wildlife
endocrinology suggest that faecal glucocorticoid metabolites (fGCM) can be used to
study adrenal activity remotely, to assess the Hypothalamic-Pituitary-Adrenal (HPA) axis
response towards stressors. The aim is to assess the impact of translocation on wild Asian
elephants in Peninsular Malaysia using faecal endocrinology and GPS technology. The
specific objectives are: (i) adapting hormone sampling methods for use under tropical
field conditions, (ii) comparing fGCM concentrations between translocated and local
resident elephants using enzyme immunoassay
● Findings:
- Translocated wild Asian elephants had lower basal fGCM profile than resident
local elephants when compared between 2 months up to a year after transclocation
- After Jalong’s(elephant’s name) translocation near a highway, she showed
repetitive movements alongside the highway but avoided crossing it, Jalong’s
initial fGCM was similar to the fGCM concentration to other translocated
elephants. Before Jalong crossed the highway for the first time, an increase in
fGCM concentration was detected, the timing coincided with Jalong’s delivery of
her calf which may have result in the in an increase in fGCM. Her hormones
remained elevated after crossing the road (this could be related to the challenges
of exploring a new environment) - this was seen as a positive indication that she
was actively coping with challenges in her surroundings & caring for her young

Behaviour Sampling Techniques

Measuring Behaviour:

1. Events (instantaneous) vs states (durations)

 ● Behaviour may be regarded as events or states. If we consider behaviour at the moment
of their onset/at any defining instant = recording events
● Events are short-duration actions that begin and end quickly
● States are longer duration processes
● For example if we record that an animal assumes a sitting posture, an act that occurs at an
instant (event)/ if that animal is seated (state)
● For event; can be defined in terms of frequency of behaviour : (i) total frequency (all
occurence of behaviours of interest is recorded), (ii) partial frequencies ( iii) the fact the
behaviour occured at least once or zero (one-zero sampling)
● For states; can be measured by duration

Constructing An Ethogram

● When observing animal behavior, scientists maintain a record by constructing an

ethogram
● An ethogram contains predetermined descriptions of expected behaviors
● Contains both events as well as states
● As well as a description of what the behavior entails
● E.g. sleeping (a state) – described as lying down with eyes closed
● Paw swipe (an event) – described as fast movement of paw towards other individual
● Can contain inferred consequences: e.g. exploring
In order to construct a record of animal behavior, you need to first figure out how you are going
to sample animal behavior
Turns out that there are a number of methods

Methods for behaviour observation

Ad libtium ● opportunistic sampling of a certain behavior. kind of just

write down whatever is occurring.
● good for experiencing rare behaviors and preliminary data
● bad because it is bias toward age groups and misses brief
instances of behavior
● Aim is to capture every behavioral event that occurs while
sampling for all animals
● Done by writing as fast as you can what you’re observing
● Usually conducted with the help of video cameras
● Behaviors are meticulously noted down in the lab afterwards by
reviewing recorded videos
● No rules on timing & individuals. Non systematic. Informal
observations. Biased towards recording conspicuous
individual/behaviour

Focal sampling ● observing a certain individual for a certain period of time

 (all occurence or specified actions of an individual of specified group). A
record is made of the length of each sample period and for each focal
individual
● Can rotate observations after that
● Good: records events and states
● helps define sequences of behaviors
● bad because sometimes don't see animal and the observed
behavior isn't typical. hard in the field

Instantaneous scan ● A very common method of sampling behavior

sampling ● Rather than sampling continuously, the observer samples
behavior at set intervals over a period
● E.g. sample a behavior once every minute for one hour
● Noting down 60 behaviors for each animal observed
● ¨Is scan sampling more appropriate for accurately measuring
events or states?
● Events are short-duration processes while states are long-duration
processes
● Recording only for 1 second out of every 60 seconds

One-Zero ● For a definite short periods of observation time, the recording of

Sampling occurrence/ non-occurrence of certain pre-defined behaviour

Sampling Errors

Clever Hans effect describe when an animal or a person senses what someone
wants them to do, even although they are not deliberately
being given signals.

Observer's effect Presence of the observer changes behaviour of the animal

Observer’s biases Unstandardized observations. Biased towards recording conspicuous

individuals/behaviour

Observer’s fatigue Over time, observer gets tired and makes less accurate observations
Tagging effect Tags may effect behaviour of an animal