animal design is affected by previous lineages (eg. rabbits can't produce birds), laws of physics., and trade-offs (eg. body size and flight maneuverability in
birds)
Darwinian evolution
living individuals are descendants of other individuals that lived in the past
evolution occurs through cumulative change over a long period of time
new forms occur via splitting of
lineages
natural selection is an engine of
change
challenging the model
genealogies
fossils
living forms
biogeography
differences between organisms should reflect their niches
eg. comparative approach with African weaver birds
individuals should possess advantageous characteristics for solving day-to-day problems
eg. optimality (utility)
theory
eg. evolutionary game theory
Studying behaviour scientifically
behaviour: the measurable response of an organism to stimuli from its environment
how can this be quantified?
why study animal behaviour?
links physiology and morphology of an organism to its environment
essential for reproduction, therefore under strong natural selection
ideal for studying evolutionary mechanisms
crucial for effective conservation in natural and captive situations
important for production of domestic animals and training of companion animals
provides insights into human health and behaviour
history of studying behaviour
Darwin
based studies on description and observation
behaviourists (eg. Thorndike, Watson, Skinner)
based studies on controlled laboratory experiments on a small range of animal;s
ethologists (eg. Lorenz, von Frisch, Tinbergen)
based studies on observations in the natural world on a range of animals
pyschology
behaviourism, similar to the work of behaviourists
comparative, study different animals in order to understand humans
eg. studies of animal intelligence
ethology
behavioural ecology
applied
for purposes of welfare research and animal science
studious
behavioural biology (present)
Tinbergen's 4 whys
proximate
explanations for the existence of behaviours over an animal's lifetime
development (ontogeny)
how did the behaviour develop in the organism's lifetime?
eg. learned, product of sexual maturity
mechanism (causation)
why is the organism exhibiting the behaviour right now?
ultimate
explanations for the existence of behaviours over a lineage (over evolutionary time)
evolution (phylogeny)
where/how did this behaviour develop over time?
function (adaptation)
what value does the behaviour provide to the fitness of the lineage as a whole?
adaptation: a trait that increases the fitness of the animal compared to its alternatives
"design"
confusions in evolutionary theory
sex ratios in animal populations
infanticide in primates
incest avoidance
Primate diversity and ecology
why do we study primates?
reasoning by homology (phylogenetic approach)
humans ARE primates and share similarities due to common descent
reasoning by analogy (functional approach)
primates provide models for how ecological pressures shaped adaptations in environments occupied by human ancestors
primates are diverse in body size, habitat, diet, social organisation, and activity patterns - how did evolution shape these behaviours?
shared characteristics between primates and humans
grasping hands
developed vision
long juvenile periods
relatively large brain
physiological and cognitive structures
behaviour
what is a primate?
characteristics:
grasping hands and feet
nails instead of claws
hind-limb-driven locomotion
reduced olfaction, enhanced vision
forward-facing eyes encased in bone
relatively large brain
long gestation
small litters (one or two)
long juvenile period
long lifespan
increased dependence on learning and behavioural flexibility
primate biogeography
mostly forest-dwelling, arboreal animals in tropical areas
modern range: Americas, Africa, Asia
ancient range: North America, Europe
primate taxonomy
about 300 different species of primates in the world
not all equally related to one another
can be split into two suborders based on genetic and morphological data
strepsirrhines
wet-nosed (do not have an upper lip)
haplorhines
dry-nosed (have an upper lip)
primate phylogeny
Lemurs
used to be very diverse, but many species are now extinct
now exist mainly in Madagascar
tend to have female-dominant societies
diverse in size and locomotion
Lorises (lorisoformes)
Africa and Asia
split into galagos (bush babies) and lorises
small, arboreal, nocturnal
Tarsius (tarsiformes)
Asia
small, nocturnal, insect-eaters
vertical clingers and leapers
New World monkeys (Platyrrhines)
south and central America
FINISH FOR REVISION
primate ecology
how do primates survive?
how do they find enough food to survive and reproduce?
how do they avoid becoming food for predators?
what is the relationship between ecology, social organisation, and behaviour in primates?
primate diets
how much food is required per unit of time affects how the day is organised
how much food is required is affected by:
basal metabolic rate: the rate at which the body uses energy while at rest to maintain vital functions such as breathing and keeping warm
active metabolic rate: the rate at which the body uses energy while active, such as when hunting or grooming
growth rate
reproductive effort
primates require:
carbohydrates (produced by plants)
amino acids (proteins)
fats and oils (from seeds, insects, and animal prey)
vitamins and minerals
water
primates must avoid toxins such as those found in adult leaves
insectivores
small body size
sharp cusps on teeth
high, sharp crests of molar teeth
simple digestive system
folivores
large body size
small incisors
sharp, shearing crests on molars
enlarged, well-developed digestive systems
frugivores
medium body size
large, broad incisors
low-cusped, relatively flat molars
relatively large digestive systems, but not specialised like that found in folivores
gumnivore
relatively small body size
long, robust incisors to tap directly into the phloem of plants
some have claws (Callitrichids)
diets and home ranges
how does the diet of a primate affect its home range?
availability of food
leaves and seeds are often more plentiful than fruit
seasonality of food
ripe fruit and leaves are not always present
primate territoriality
some primates are quite territorial
home ranges do not overlap, and territories are defended aggressively
territorial in order to defend mates (primarily males) and defend resources (primarily females)
some primates are not territorial
home ranges overlap
whether a primate is territorial or not depends on the cost of defending an area and the benefits gained by protecting limited resources/mates
predation
threats from:
snakes
big cats
birds of prey
crocodiles and alligators
defenses against predation
interspecific associations, eg. between deer and primates
small, terrestrial primates are the most susceptible
in areas where many felids or raptors exist, primates become more arboreal (eg. in South American forests), and in areas with few felids and raptors, primates evolve larger body size and group size (eg. Asian forests)
vocalisations (eg. vervet monkeys)
increased group size
detection: more eyes to spot a predator
deference: mobbing behaviour
dilution: individual risk of death is lower
primate sociality
why are primates social?
predator avoidance
feeding competition
can defend food patches
female distribution is related to resource distribution = female social groups
can be costly as there is more competition for food and mates, and disease transmission is more of a risk
social groups
solitary
polygynous (one male, multiple females)
pair-bonded
polyandrous (multiple males, one female)
polygynandrous (multiple males, multiple females)
Primate mating systems
mating systems: the ways animals find mates and care for offspring
basic rules for mammals (though there are some different variations in primates)
sexual reproduction
female gestation
female lactation
language of adaptive explanations
strategy: evolved behaviours that are the product of natural selection
costs and benefits: trade-offs in reproductive success
evolution of reproductive strategies: female
variation in parental care in animals (eg. sea horses, frogs, birds, mammals)
in mammals, female care is critical and male care varies
evolution of reproductive strategies: male
trade-offs
males can put energy into finding new mates if finding mates is relatively easy and if offspring can be raised by one parent, or put energy into helping offspring survive
reproductive strategies: female
female primates invest heavily in offspring
gestation
long pregnancies
brain growth is energetically expensive
lactation
long infant periods
energetically expensive (can cost up to 8x basal metabolic rate)
very few offspring
quality over quantity
what influences female reproductive success?
resource abundance
more food = more offspring (shown in macaque provisioning studies)
maturity
young mothers have to allocate energy towards their own growth
50% higher infant mortality rate
lack of experience
older mothers may be undergoing senescence (the process of deterioration of the body with age) so may not be fit to carry offspring
longevity (how long an individual lives)
the longer a primate lives, the more opportunities it has to produce offspring)
accounts for 50-70% of variance
group size
smaller groups tend to have higher reproductive success
rank
higher ranked females tend to have more offspring
higher ranking females get greater access to resources
rank is determined using a dominance matrix to identify relationships of dominance within groups
sociality
females with more social bonds produce more offspring
eg. female baboons with strong social bonds have greater reproductive success, and live longer
reproductive trade-offs
resources are limited
allocation of energy to one offspring comes at the expense of others
there is a trade-off between quantity and quality of offspring
eg. lactational amennorhea prevents pregnancy while a female is lactating
weaning allows females to get pregnant again
weaning
infants become more independent
nursing is gradually reduced
energy becomes available for the mother to conceive again
time of weaning varies in different primates
sexual selection and male mating strategies
limiting factors for reproductive success
in females: adequate resources
in males: access to females
sexual dimorphism: distinct differences in size or appearances between the two sexes
eg. the tail feathers of a male peacock
sexual selection
because of high sexual selection in some species, a lot of resources go towards increasing a male's mating success rather than going to females to increase reproductive success
arguably stronger than natural selection, but both may balance out over time
eg. large tail feathers in peacocks = difficulty flying = more likely to be eaten by predators, so only peacocks with tail feathers below a certain range can survive to breed
male reproductive success is more variable than females
intrasexual selection
male competition
eg. stags during the rut
dominant males gain access to females
intersexual selection
female choice
the female selects the most attractive male
sexual dimorphism and mating systems
pair-bonds have little male-male competition, so there is little dimorphism
polygynous groups have huge body and canine dimorphism
polygynandrous groups have large dimorphism
sperm competition and mating systems
polygynandrous groups
estrus females mate with many males, therefore males with larger sperm volume have a greater chance of fathering offspring
selects for testes size
investing males
pair-bonding
mate guarding (mates may "cheat"), eg. by grooming often
increased parental investment in offspring
cooperative breeding
males and other individuals in the group help to raise the young
eg. marmosets and tamarins often have twins, so the mother can increase fertility/survival rate by having helpers
humans are cooperative breeders
male-male competiton
behavioural flexibility in primates
in polygynandrous groups, males establish a dominance hierarchy
higher ranking individuals have greater reproductive success
infanticide
death of an infant accelerates the return of a female to sexual receptivity/lactational amenorrhea)
85% of deaths follow takeover by a new male
unweaned infants primarily targeted
rarely done by sexually active males in the group
in 45-70% of cases, males mated with the same female
infanticide is not reproductively beneficial for females, therefore they have evolved counterstrategies
paternity confusion
male friendships
male protects female in return for grooming
studies show that female stress rises if the female does not have a male friend during a takeover
naturalistic fallacy
what we see in nature is not necessarily "correct"
explanations do not equal justifications
Primates to Hominins
origin and evolution of mammals
early Triassic period
Therapsids
225 million years ago
overlapped with dinosaurs
warm-blooded, fur-covered reptiles
egg-layers, did not produce milk
around for 30-40 million years
late Triassic, early Jurassic period
first mammals
dinosaurs ruled
oviparous (laid eggs)
end of Cretaceous
dinosaurs extinct, increase of mammal radiation into new niches
origin of primates
angiosperm hypothesis
the adaptive radiation of primates occurred with the radiation of angiosperms (flowering plants) that offered new opportunities and an unexplored niche
omnivores adapted more flexible hands to handle fruit
visual predation hypothesis
omnivores adapted orbital convergence (eyes moved from sides of head to front) in order to forage for fruit nocturnally
leaping hypothesis
primate biomechanics for leaping and grasping were favoured
generalised nocturnal characteristics (large forward-facing eyes, grasping hands and feet) were favoured
terminal branch hypothesis
evolved to better meet the needs of an arboreal lifestyle (vision, leaping, grasping hands and feet)
evolution of early primates
Plesiadapiformes
frugivore
no opposable thumbs
nails on one toe, claws on the others
long, narrow snout
small brain
large incisors
Eocene period
earth was warm and wet
tropical forests spread into North America and Europe
evolution of two types of primates (it is not possible to say which came first)
Adapids
possible Strepsirrhine ancestor (wet-nosed primate)
larger than Omomyids
diurnal and quadrupedal
Omomyids
possible Haplorhine ancestor (dry-nosed primate)
possible tarsier ancestor (even though tarsiers are strepsirrhines)
nocturnal
some leapers
evolution of Haplorhines and Anthropoids
evolution in Fayum, Egypt during the Eocene/Oligocene boundary (33-36 million years ago)