Primates and Primatology Flashcards
Name the distinctive features of mammals
- hair- fur
- sweat glands
- mammary glands (milk production)
- 3 middle ear bones
- specialised teeth
- 4 chamber heart
- neocortex in brain (outer layer involved with cognitive ability)
Name the characteristics of primates
- clavicle (allows for arm movements/living in trees, locomotion)
- opposable thumbs (ability to grasp, precision grip)
- fingernails
- binocular and colour vision
- generalised dentition (allows omnivore diet)
- slow reproductive rates (lower fertility rates than expected for size)
- larger brains than expected for body weight
Name different classes of primates
- prosimians
- Tarsiers
- Platyrrhines (New world monkeys)
- cercopithecines
- colobines
- Apes
Describe Madagascan Prosimians (including examples
Examples:
- Brown ruffed Lemue (Varecia rubra)
- Ring-tailed Lemur (Lemur catta)
- Aye-aye (Daubentonia madagascarensis)
- Mouse Lemur (Lemur pusillus)
Characteristics:
- small size
- eat insects/fruit
- mainly solitary
- most ancient
- rely on smell so have snout/muzzle
- specialised glands
- Aye Aye are solitary
- in Lemurs, females are dominant
- most are nocturnal
- Males home-ranges larger than female‘s
Describe African and Asian Prosimians
Examples:
- Angwantibo (Arctocebus spp.)
- Lorises (Loris spp.)
- Potto (Perodicticus spp.)
- Bushbabies (Galago spp.)
Describe Tarsiers
- south eastAsoa
- endangered
Describe Platyrrhines
- South and Central America
- members of the group help to raise offspring of dominant couple
- see world in 2 colours
Examples:
- marmosets
- tamarins (saguinus spp.)
- Capuchins (Cebus spp.)
Characteristics:
- Variable body size
- Dichromatic vision
- Group-living- often cooperative breeders
- Mostly diurnal
- Variable social systems (harems, multi-male multi-female)
- Capuchins: „chimpanzees of America“
Describe Cercopithecines
- Africa and Asia
- varied diet
- multiple males and females in groups
- multi-chambered stomach
Examples:
- Baboons (papio spp.)
- Mandrills (Mandrillus sphinx)
- Macaques (Macaca spp.)
Describe Colobines
- Africa and Asia
- 1 male unit
- Infanticide occurs
Examples:
- colobes
- leaf-momekeys
- snub-nosed monkeys
Describe Apes
- larger brain
- live during day
Examples:
- Gibbons
Orang-utangs
- gorillas
- chimpanzees
- Bonobos
- Humans
Discuss different ways of studying primates
- Observation vs experiments
- Field vs captivity
Name advantages and disadvantages of studying primates in capivity
- advantage- can con tool extraneous variables such as diet and interaction
- disadvantage- behaviour will not necessarily be ecological relevant
What is adaptation
- characteristics that improve an organism‘s chances for survival and/or reproduction
- functional traits passed down through the next generations
- maintained through evolution and selection
Describe the process of selection
- variation present amongst individuals (due to genes)
- selective pressures- affect ability to survive and reproduce e.g.:
- biotic environment (climate/habitat/substrate),
- biotic environment (food, predators, diseases, other species, attraction, competition- intraspecific)
- some individuals survive and others don’t
What are the mechanisms of selection in natural vs sexual selection
- natural- survival and reproduction
- sexual- reproduction
outline genetic drift
- bottleneck- only few individuals survive- distribution of characteristics change
What are 3 factors needed for adaptation/selection
- variation in traits
- heritability of traits
- selective pressures
Name different domains of primate adaptations/functional traits and the selective pressure that causes the,
- limbs- habitat
- teeth- diet
- sense/brain- food/diurnal life/ sociality
- life history (birth, death etc)- large brain 9developmental and energetic constraints)
What are homologous chatracteristics
Present in a common ancestor
What are analogous characteristics
- present in some apes/primates (not common ancestor)
- caused by convergent evolution (e.g. birds and bats both developed wings)
- evolved later in humans
- aka homoplasy
What does the comparative approach allow
- Identification of correlated factors
- Build evolutionary theories e.g. look at skull shape from fossil evidence- can understand evolutionary history e.g. when specific organisms walked
- Infer selective processes (e.g. how behaviour is shaped by social and environmental conditions)
- Infer events of convergent evolution
Outline primate limbs and locomotion
- hand/feet- high prehensility
- 5 digits
- opposable thumb- allows hook to swing from branch ti branch, also use tail
- divergent and partially posable big toe
- nails instead of Claws
- tactile pads with enriched sensory nerves of digits tips- precision of manipulation of objects
–> 55 million years ago then lineage of primates emerged, had to rely more on catching preys- had to find fruit in trees - presence of clavicle- mobile arms and shoulders- pre-adaptation for tool use
Outline differing locomotion in primates
- clinging/leaping e.g. galago
- quadrupedalism e.g. lemurs
- brachiation- hangers e.g. spider monkeys
- bipedalism
Outline the comparative approach in terms of primate limbs and locomotion
- can compare skeletons e.f. of arboreal and terrestrial- terrestrial has stretched upper limbs whereas arboreal is using low strides- different behaviour results in morphological differences e.g. relative length of bones
- can compare hand shapes and bones etc- link between bone structure and locomotion- can infer locomotion of evolutionary ancestors
Describe primate diet and teeth
- generalised dentition- adapted to diverse diet- lack of specialisation
- baboons and chimpanzees omnovorse- leaves, seeds etc
- some more specialisations:
- cercopithecines- frugivores (fruits)
- colobines/gorlias- folivores (leaves/grass)- specialised multiplee chambers
- some prosimians- insectivores (arthropods)
- due to changes in gut and teeth
Describe the comparative approach in relation to diet and teeth
- distantly related primates- similar diet associated with certain tooth form
- similarity in characteristics corresponds to convergent evolution (homoplacy) and adaptive characteristics
Describe variationn in primate teeth which isn’t to do with diet
- sexual dimporphism- occur differently in male and females of same species e.g. short vs long k9s in makkaks
- different teeth length due to competition
Describe primates neural adaptations
- allometric scaling- compare brain size relative to body size- most allocated above allometric line
- general law is that correlates with body size, adaptation is when above/below
Describe primates sensory adaptations
- Rely on vision rather than olfaction- reduced snout, flat nose, reduction of olfaction brain structures
- morphological adaptations- skull, eyes, brain
- Colour vision (2/3)
- stereotypic vision (3D, right nd left visual fields cross)- allows to live in trees, catch insects to side
- forward facing eyes
describe the primate life history
Lite hosiery is timing and pace of major life events:
gestation - birth- growth- maturation- reproduction- senescence- death (no menopause)
describe adaptation in the primate life history
- slow reproductive rates and fertility rates greater longevity
- large brains causes slow maturation and growth, which therefore causes slow life history
describe consequences of the slow life history in primates
- Intense parental/maternal care
- Great dependence on flexible learned behaviour e.g. orangoutangs from mothers
- Life in social groups
- Mostly diurnal
- Stable groups
- Long-term relationships
- Cooperation
- Overlapping generations
- Social learning, cultures
outline biological classification
- taxonomy- description, identification, nomenclature and classification of organisms
- Carl Linnaeus- 18th century
- based on common characteristics
- DMPCOFGS
outline species
- 180-300 primate species
- definitions- either cant reproduce (biological barrier), cant meet (geographic barrier), sometimes unclear- hybrids e.g. humans and neadrathals
Describe Cercopithecines and Colobines characteristics
- Variable body size
- Group-living
- Hierarchical dominance
- Female philopatry
- Variable social systems
(harems, multi-male multi-female) - Strictly diurnal
- Sexual dimorphism
- Colobines: folivores, „cow“ stomach, harem systems, infanticide common
-Cercopithecines: frugivores/omnivores, cheek pouches, multi-male multi- female, female sexual swellings (not in guenons)
describe characteristics of lesser apes
- Gibbons and Siamangs
- No tail Monogamous
- Duet song
- Male and female dispersal
- Strictly diurnal
- Brachiation
ape characteristics
List the requirements of a primates diet
Provides energy for growth and essential elements for locomotion and nutrition
- carbohydrates- sugar
- amino acids
- fat/oil- energy storage
- vitamins/minerals
- water
- AVOID- toxins incl. tannins, caffein, alkaloids
Where do primates obtain required nutritional elements from
- carbohydrates- fruits, seeds, gum
- proteins- young leaves, arthropods, animal preys
- fat and oil- seeds, animal preys
- vitamins and minerals- fruits, animal preys, geophagy (earth elements e.g. clay/rocks)
- water- food, tree holes, water holes
Describe how primates can get access to water from tree holes
- tool use- fluid dipping- wooden sticks- chew end (used by chimpanzees), also used for honey
- leaf sponges
outline other feeding strategies in primates
- sits on branches and pushes ground to get termites and ants
- nut cracking- pounding techniques, Woden hammer
- tortoise pounding- Catcha and kill tortoises to extract meat
- oyster cracking in long tail macaques
–> use of social learning
–> similar techniques across species
Outline the influence of colour and stereotypic vision on feeding strategies
- some can see in 2/3 colours
- colour vision allows to see varying degrees of brightness of fruits
- can see in 3D- can capture moving oral and move in trees
Outline the influence of colour and stereotypic vision on feeding strategies
- shape of teeth adapted to certain diets
- longer dinger in aye-aye
Outline frugivory (what it is, competeition, issues)
- eat mostly fruits
- fruits are source of sugars and carbohydrates
- resources worth fighting for
- issues- patchy and seasonal distribution/production, Hugh unpredictability, monopolisable, variation between years as well as within
- causes contest completion due to clumped resources- resources are scarce and valuable
- vertical distribution of fruits and ripeness- causes vertical contest competition- best fruits at top (as sunlight)- female chimpanzees that are dominant can have highest reach in trees
Outline folivory
- eat leaves and grass
- provides source of fibres and proteins
- scramble competition as resources evenly dispersed- once fid, eat as much as can
- resources not worth fighting over, no direct competition
- hard to monopolise
- issues- secondary components, hard to digest, toxic- so developed stomach with multiple chambers
Outline insectivory (what it is, competition, issues)
- source of proteins and fat
- moving preys- harder to catch
- scramble competition
- means occupy different heights- niche partitioning
Outline movements influenced by food availability and distribution
- depending on how much food availability, have different travelling distance
- travel less during day in times of low food availability
Outline fission-fusion (troops) influenced by food availability and distribution
Chimpanzees:
- party size changes
- when low food, party size decreases
Outline resource defence influenced by food availability and distribution
- core area defence- more food in core
- food patch defence- no difference between food in core and periphery
- boundary defence- more food in periphery
- size of core vs periphery area and different stability of resources affects formation
Outline spatial memory effected by food acquisition
- primates need too remember location of feeding sites so can return to certain areas ay certain seasons
- can map routes taken by primates- if know location, shoudlnrtravel in direct line at greater speed, rather than trial and error
Outline niche partitioning
- partition selves
- differ in food specialisation
- sympatric species
- allows to form large groups of multiple species- interspecific associations
outline anticipation in primates
- e.g. anticipate eating figs at first time of day
- leave earlier to site in order to be first at trees- leave first for fogs as most sought after
Outline interspecific associations
- improves foraging efficiency as can occupy strata in first that dent usually occupy
- increase of arthropod intake when in association- broader niche
name predators of primates
- snakes
- birds
- cats
- humans
- fossa
- Komodo dragons
- dogs
name anti predation strategies
- grouping- dilution effect (Hugh number of potential targets), confusion effects (leave in many different directions)- interspecific association
- sleeping sites- nests building (apes), cliff sleeping sites (lemurs, geladas)
- alarm call systems- inform other animals and others in group
Responses cam be differential e.g. for leopards use referential alarm calls and climb trees, whereas for eagles use referential alarm calls and go down from trees
alarm call systems graphs
type of calls different for each type of predator
Outline the co-evolution principle (ecological importance of primates)
- Reciprocal evolutionary changes between pairs of species or groups as they interact with one another
- angiosperm co-evolution hypothesis-our primate ancestors evolved key adaptations like forward-facing eyes, excellent colour vision, rounded, blunt teeth and fingers without claws, all for the purpose of eating and living from fruits
outline the visual predation hypothesis
forward-facing eyes and grasping hands evolved in primates to allow them to eat insects in trees
describe other ecological importances of primates
- seed dispersal- pre dispersal, dispersal, and post dispersal
- high percentage of seeds are not desyroyed- all primate wither spit or swallow seeds away from original tree- causes tree growth in different areas
- prefer to groom in open areas away from trees- causes greater dispersal
- NWM + OWM specialized in colourful fruits with flesh with seeds easy to swallow
- High number of seeds and species dispersed
- High number of seeds dispersed over long distances
Give details of the life history of primates (including time periods)
- late maturation –> long juvenile period –> long maternal investment (lactation, carrying, protection)
- Slow reproduction: average inter-birth interval 2.3 years (max 7-8 years orang-utans)
- Single offspring (max. 2 Callitrichids)
- trade of between longer life and limited number of offspring- invest energy into current offspring rather than next generation
- low adult mortality, long reproductive career, long life span, females outlive males
- Little Mama- died at 79
What is sexual selection
- Selective process acting on behaviour and traits that maximise reproduction (form of natural selection with focus on reproduction rather than survival)
- Trade-off between natural selection and sexual selection adaptive value of traits
Describe intersexual vs intrasexual selection
- InTRA- intimidating, deterring or defeating same sex rivals (competition for sex between members of same sex)
- inTER- making themselves attractive to the opposite sex (mate choice), opposite sex reproductive strategies differ (and compete)- sexual conflict
What do all reproductive strategies do
Maximise lifetime reproductive success
Outline gender-differentiated patterns of sexual selection in males and females
- males are ready to mate most of the time, whereas females are constrained by physiology (gestation/lactation) and maternal care
- thus means there is a limited humber of receptive females
- means that males compete yo access females (male-male competition), and females choose among potential mates (mate choice)
Describe sexual dimorphism in terms of intrasexual selection
-male-male sexual competition
- males have larger body size
- use physical weaponry e..g size of k9
- vocal apparatus
- secondary sexual characteristics e.g. flanges in orangoutangs- develop after sexual maturity- depends on food availability
- body mass- relationship to mating systems in primates
- Higher sexual dimorphism in mating systems with one single reproductive male
Outline intrasexual competition in terms of sperm competition
- male-male sexual competition
- relates to mating systems
- sperm compete in female genital tract
- reflected in tests size
- chimpanzees large testes whereas gorillas small (as live in group with 1 reproductive male)
- High sperm competition in multi-male societies
Outline intersexual competition in terms of mate choice
- Mate selection depends on attractiveness of certain traits
- Traits related to resource holding potential (RHP)- capacity to provide protection, parental care, resources, good genes
- RHP (often) given by honest signals (by males)- good genes- means healthy individual - ornaments in mandrills, douc lanurus and uakaris
- e.g. brightly coloured male mandrils get more copulations- females approach more brightly coloured males, and present more to brightly coloured males- indirect indicator of good health
Outline intersexual selection in primates in terms of sexual conflicts
- interests of males and females differ
- Males and females prefer different reproductive strategies to reach
optimal fitness - Conflicting optimal fitness strategies concerning reproduction
- Evolutionary arm race between males and females
Outline intersexual selection in terms of reproductive strategies
- male reproduction cheap- invest in numerous gametes with no energetic reserved- ready to mate most of time
- female costly- invest in few gametes with high energetic reserves, invest in costly maternal investment (gestation, lactation, maternal care)
- means for females reproductive mistakes are costly, makes females choosy, and means that males are limited by access to females
Describe the implications of gender differing reproductive strategies on space use and competition
- females- high investment so limited by the access to resources (food)
- success determined by number of offspring they can produce
- means compete among females for better resources (shorter inter-birth interval, higher survival, better offspring survival, better health)- causes to monopolise resources, dominance hierarchies and matrilines (dominance hierarchies passed onto daughters)
- male- low investment- soserachc for more matings- so limited by access to females
Outline dominance hierarchies in females
- separate from hierarchies in males
- indirect fitness advantages- access to food resources- higher rank forage more without moving (better physical condition for pregnancy)
- direct fitness advantages- survival- better condition, reduced vulnerability to predation
- faster reproduction (as shorter inter birth interval)
Outline dominance hierarchies in males
- reproductive sucess- leads to high variance in reproductive success
- higher ranking males get more mating opportunities
- means get high reproductive skew- limited number of rales get most offspring
- when more males, less have change to mate
- Maxaques- males in prime sire the most offspring- top sire in each year produced 24% of the infants, while 71% of troop males sired no offspring at all
Outline sexual coercion (male reproductive strategies)
- form of sexual conflict as done without consent of females
- males attempt to bias mating in their favour by using harassment, intimidation and forced matings
- leads to higher rate of potentially fertile matings, a lower rate for other potential mates, and costs for the females
- cost for females so have developed counterstrategies- Costlier investment from females to counteract male sexual coercion
outline female counterstrategies to male coercion
- female coalitions
- female dominance
- “When females formed coalitions, 100% won against a target male(s)”
- all targets of female coalitions were males, never to attack other males
Outline infanticide as a male reproductive strategy
- will commit infanticide if now they are not the father
- Faster reproduction- species with lactational amenorrhea (infertility after giving birth due to lactation), short male tenure
- Infants of other males targeted- New males should do it, no prior mating with females
- High chance of siring new infants Dominant males, one-male groups
Outline female counter strategies to infanticide
- promiscuous mating- paternity confusion (means men not Sur if they are the father)- formation multi female and multi female social systems
- synchrony of reproductive cycles- when cycling together, can mate with men from other units- one male cant keep access- confuse paternity
- post-partum oestrus- lactation reduced, so infanticide doesn’t occur when lactation longer (no postpartum oestrus, infanticide common)
- male-female association- e.g. baboons- pair bonding - more time spent with male after birth- protection against other males, decreases after death of infant
Outline female defence as a male reproductive strategy
- Males defend access to one or more females directly
- long-term associations- monogamous bonds and harems
- dominance hierarchies in mixed sex groups, dominant males have priority of access, and short term censorship nonds
- coercion- males physically force females to
Input to social systems diagram
Outline dispersed solitary social systems
- Males‘ home-range > females‘ home-range
- Forage alone
- Social contact via vocal + olfaction (scent marking)
- Often nocturnal, sleep in trees in day
- polygynous- each male mates with several females
- prosimians and orangutangs
Outline pair bonded social systems
- one male one female
- teritorial defence
- monogamous (sometimes extra-pair copulations)
- paternal offspring care
- Titi monkeys, owl monkeys, some callitrichids (marmosets, tamarins), Hylobatids (gibbons and siamangs)
Outline one female multi male social systems
- Cooperative breeding
- Reproduction of subordinate females (daughters) suppressed by aggression or pheromonal signals
- Polyandrous- more than one mating male- cooperative polyandry
- Marmosets, tamarins
Outline one male social systems (Harem)
- Single reproductive adult male defends a territory with several females and subordinate males
- Exclusive mating access
- Polygynous
- Bachelor groups
- High risk of infanticide
- Sexual dimorphism due to high male-male competition
- Colobes, langurs, guenons, patas, howler monkeys, gorillas
Outline multi male multi female social systems
- Polygynandrous
- Males and females mate with several members of the oposite sex
- Female philopatry
- Matrilines
- Exaggerated sexual swellings common
- Macaques, most baboons, vervet monkeys, capuchins, spider monkeys, wooly monkeys, some langurs and some lemurs
Outline fission- fusion communities
- Temporary sub-groupings depending on food conditions
- Male philopatry
- Female dispersal
- spider monkeys, colobus, chimpanzees, baboons
Outline multi-level societies
Three-levels of social structure:
- one male units- reproductive (one male, multi female)
- bands- ecological (forage together)
- troop temporary aggregations
- polygynous
- infanticide
- Hamadryas baboons, geladas, sub-nosed monkeys
Primate mating systems diagram
List impacts on primate social systems
- social structure (size and composition)
- mating system
- social organisation (interactions, space-use, competition, dispersal, philopatry)
Outline costs of primate sociality
- Competition for food
- Competition for mates
- Competition for space
- Disease transmission
- Infanticide
Outline benefits of primate sociality
- searching for food
- reduced predation risks
- competition with other groups
- mating opportunities
- friendships- social bonding
- social learning opportunities
Outline the heuristic approach to primate socio-ecology
- aim at explaining all forms of primate social organisations
- Models stemmed from natural history, improved by concepts of inclusive fitness (when harmful for self but benefit for genetic relatives), selfish herd (putting another before predator) and parental investment
- Causal relationships between food distribution, predation, mating strategies, social organisations and relationships
Outline comparative primate socio-ecology
Van Schaik, 1989:
- Multi-females groups initially favoured by predation
- Food competition determines female social relationships within and between groups
- competition regimes- contest vs scramble
Outline contest competition regimes
- clumped resources
- resources scarce and valuable
- resources worth fighting over
- resources are monopolizable
- direct competition
Outline scramble competition regimes
- dispersed (insects) or evenly (leaves) ditributed resources
- scramble to get enough
- resources are not monopolizable
- resources not worth fighting over
- no direct competition
Outline female relationships in primate socio-ecology
Van Schaik, 1989:
- consequence of two types of competition (scramble and contest), and whether within or between group competition
- scramble competition within group- females develop individualistic, egalitarian dominance hierarchies
(Type A)
- when high predation pressure, scramble competition within group and contest competition between group creates philopatric females and egalitarian hierarchies (type C)
- contest competition within groups means females develop nepotistic, despotic dominance hierarchies (type B)
- contest competition between-group when low predation pressure and high population density creates nepotistic, egalitarian hierarchies (type D)
outline ecological model of private socio-ecology including the influence of infanticide avoidance
- when low predation, high risk of infanticide
- female dispersal and association with resident males
- Sterck et al, 1997- new labels to van Schaik model
- type A- dispersal-egalitarian
- type B- resident nepotistic
- Type C- resident-egalitarian
- type D– resident-nepotistic-tolerant
Outline factors that contribute to female social relationships in primates
Sterck et al, 1997:
- food distribution and predation risk effects female gregariousness
- infanticide risk effects how females associate with males, which may lead to females sharing the same protector male, which also influence gregariousness
- gregariousness effects attachment of males, ad creates a competitive regime (ethic is also affected by habitat saturation)
- this creates social relationships
Outline an example of the impact of varying levels of within-group contest competition on primate socio-ecology (across species and sites)
Across species- Mitchell et al, 1991:
- Saimiri oerstedi of Costa Rica (high WGCC) compared to Saimiri sciureus of Peru (low WGCC)- similar predation level and group size
- higher WGCC lead to linear dominance hierarchies and female philopatry
- low WGCC lead to a lack of linear dominance hierarchies, and creation of female dispersal
Across sites- Koeing et al, 1998:
- presbytis entellus of southern Nepal and India across two sites
- same patterns as above found for high vs low WGCC across the sites
Outline the link between scramble competition within group and group size
- should support large group sizes, but group sizes relatively small in reality
- may be due to folivore paradox- patch depletion may limit group size expansion
- female reproductive success may be better in intermediate group sizes
- females may disperse to avoid male harassment/infanticide
Outline examples of inconsistencies in primate socio-ecology
- Saj et al, 2007- Colobus vellerosus- folivore- more F-F affiliation than F-M affiliation, demale dispersal, no female coalitions over food and low F-F grooming- suggests dispersal-egalitarian- may be response to risk of infanticide
- chimpanzees and bonobos- frugivores- female philopatry predicted,, but find male philopatry; no female hierarchy expected, but see linear female dominance hierarchy and bonobo female coalitions
Outline potential additional factors to the socio-ecological model in primates
- role of phylogenetic constraints- some traits inherited and so conserved independent of ecological conditions
- role of male mating strategies on female-female competition?
- constraints on group size enforce female dispersal?
- demography alters rates of alliances and aggression?
- quantification of predation pressure and competition- lack of method standardisation
Outline the links between predation and female gregariousness
- De Ruiter, 1986- wedge-capped capuchins- predation benefits of group living- allows more scanning
- increased time spent in vigilance when in small groups
- Large groups occupy lower strata, small groups occupy higher strata
Outline the link between infanticide risk and female gregariousness in primates
- leads to association with likely father
- mountain gorillas- female transfer subsequent or simultaneous to infanticide- females join the infanticidal male (Watts, 1989)
Strengths and limitations of the primate socio-ecological model
Strengths:
- appropriate model to explain primate social systems
- food distribution, predation and infanticide risk as main determinants
- contest vs scramble competition
Limitations:
- Inconsistencies to relate competition, dispersal and social relationships
- folivore paradox
Outline co-operation
- Biological process where individuals or groups act together
- can be between different species where both benefit (mutualism), or between the same species where there are common benefits (group living)
- widespread phenomenon across living organisms e.g. for sociality, anti-predation, forging efficiency, hunting success, thermoregulation, territorial defence, paternal care, and cooperative breeding
Outline cooperation in relation to evolution
- evolutionary puzzle
- natural selection- favoured behaviour enhances success of individuals
- cooperation, however, mean that behaviour has cost for the cooperator
- involves alturism- Behaviour that benefits the fitness of the recipient at a fitness cost for the giver/actor- creates risk of exploitation by self-interested free-riders
List theories of evolution of cooperation
- kin selection
- reciprocal alturism
- group selection
Outline the kin selection model of cooperation
Smith, 1964; Queller, 1992:
- Cooperation with family members
- An altruistic allele can improve its prevalence in the gene pool by promoting the success of the same allele in other individuals
- Individuals increase their inclusive fitness by being altruistic to relatives, who share a portion of their genes
- Hamilton rule- altruism evolves when the costs are smaller than the benefits multiplied with degree of relatedness of benefits
- inclusive fitness involves individual own fitness, as well as fitness of each relative- weighted by the degree of relatedness
Outline the reciprocal altruism model of cooperation
Trivers, 1971:
- individuals cooperate with those that cooperate with them
- If costs of cooperation are low and benefits are high, altruistic alleles can spread by mutually promoting their success
- individuals reciprocate altruistic acts, leading to the development of long-term social relationships
Outline the group selection model of cooperation
Wilson, 1975; Smith, 1976:
- Groups of cooperators outcompete groups of non-cooperators
Outline the influence of social relationships on cooperation
- social relationships are made of dominance and friendship
- have rocks of aggression, passive benefits of tolerance near resources, and active benefits of exchange off commodities (grooming, food sharing, and antagonistic support)
- risk and benefit balance modulated by kinship, friendship and reciprocity
- proximity may be proxy for friendship
Outline the influence of grooming on social relationships
- most important way to form friendships
- other benefits: reduces stress, reduce parasitic load
- often bottom-up (give-receive)
-market commodity (grooming vs tolerance, grooming vs mating, grooming vs protection, grooming vs support)
-can be reciprocated - tolerance vs grooming in tufted capuchins (Tiddi et al, 2011)- More tolerance over food after reciprocal grooming, More grooming after being tolerant at food resources- market commodity that can be exchanged and reciprocated
Outline fitness advantages of friendships in primates
- Silk et al, 2009- influence of kinship on bonding- Wild baboons associate more with related individuals
- Females more social (with kin and non-kin) have offspring that survive better
- Silk et al, 2010- Wild baboons females with consistent stable relationships live longer
- Langergraber, 2007- Male chimpanzees- stable, long-lasting, reciprocal bonds, cooperation influenced by kinship, but also by bonds among non-related individuals
- Feldblum et al, 2021- Wild subordinate chimpanzees are more likely to sire if they have strong bonds with the alpha male
Outline food sharing in relation to cooperation
- chimpanzees- passive (tolerant) and active food sharing- active more rare
- reinforce social bonds
- link between sense of fairness and cooperation- more cooperative if have sense of fairness
Animal behaviour culture definition
behavioural practice that is shared among members of a group; is performed repeatedly over a period of time (enduring); and depends to a measurable degree on social contributions to individual learning for its appearance in new practitioners (Fragazy, 2003)
Define traditions in animal behaviour
- all group-typical behaviour patterns, shared by members of animal communities, that are to some degree reliant on socially learned and transmitted information (Laland & Hoppit, 2003)
- traditions are cultural traits (behaviours)- culture is made by the summation of them
- generalised definition including both- Behaviours that are practiced habitually in a group, rely in maintenance and transmission on social learning (McGrew, 2004)
Outline different cultiral domains in animals
- tool use
- social-sexual (greetings/gestures)
- communication (associative vocalisations, dialects)
- feeding (feeding preferences, hunting techniques)
List different ways of measuring cultural behaviour in animals
- method of exclusion
- comparing populations within site
- counting socially learned skills
List different explanations of culture in animals
- 3 factor model
- social learning
Humanities culture definition
shared patterns of behaviours and interactions, cognitive constructs and understanding that are learned by socialization- growth of a group identity fostered by social patterns unique to the group
Outline the method of exclusion (measuring cultural behaviour)
- Compare study sites
-List all behaviours
-Exclude those explained by genetic differences
-Exclude those explained by ecological differences
-Exclude universal behaviours
Outline findings about animal culture using the method of exclusion
Whiten et al, 1999:
- chimpanzee cultures in different study sites, e.g. Tai forest, Gombe
- found 39 cultural variants across social, sexual, and feeding domains
- found different functionality of behaviour e.g. some use for sexual, others for grooming
Van Schaik et al, 2003:
- orang-utang populations
- found 24 variants across social and feeding domains
- Cultural variation increases with geographic distance
- Size of cultural repertoire associated with opportunities for cultural learning- more culture if more chances for social learning
Ranger et al, 2002:
- capuchins foraging traditions
- 20 foods processed differently across sites
Outline issues with the method of exclusion for measuring animal culture
- Ecological pressures drive adaptive behaviour which can also be cultural (as socially learnt
- Genetics influence animal’s predisposition for social learning
- Animals modify their environment, also via culture (niche construction)
- Universal behaviour can also be cultural (e.g. nest-building)
- Distant populations: can’t rule out influence of genetic and ecology
Outline the method of comparing populations within site to measure cultural behaviour in animals
- compare neighbouring populations within the same site
- have same habitat, and are same sub species
Outline findings about animal culture using within site comparison
Luncz et al, 2012:
- 3 neighbouring chimpanzee communities
- nut cracking behaviour
- Stone tool use habits vary between neighbouring communities
- wooden tool use habits also vary
Luncz & Boesh, 2014:
- Nut-cracking traditions perpetuate through generations and remain
- conformity exhibited- all members of same group behave similarly, new immigrants conform to new group’s tradition
Koops, 2015:
- ant dipping tool length in Kalinzu forest (chimpanzees)
- length varies widely across chimpanzee study sites, across type of army ants and across dipping locations
- varies within the same population between two neighbouring communities- even when same type of ants and same dipping locations
Outline counting socially learned skills to measure cultural behaviour in animals
Schuppli & van Schaik, 2019:
- measures increase in general cultural repertoire
- can modify distribution of variants across behavioural domains
- reveals ecological correlates for most cultural elements
- suggests culture drives local adaptations, and is widespread and pervasive
Outline findings from counting socially learned skills to measure cultural behaviour in animals
Schuppli & van Schaik, 2019:
- orang-utans cultures- Tuanan vs Suaq
- Tuanan less sociable, and have less social learning opportunities
- greater difference between populations found than when using method of exclusion
- found ecological correlations- e.g. feeding skills necessitated by ecological conditions
- some behaviours culturally universal, but still socially learnt
Outline the 3 factor model of culture
Koop et al, 2014:
- Environment, sociality and cognition
- Considers influence of environment, sociality and cognition on culture (particularly material culture)
- Environmental opportunity, rather than necessity is main driver of material culture
- supported by Koops (2014)- ant dipping and nut cracking increases when more available, but ant still increases when more food available in general (showing matter of opportunity not necessity)
Individual vs social learning definition
- Individual- acquiring information/skills without other‘s influence- trial/error, takes time
- social- learning influenced by observation of, or interaction with, another individual or its products- fast
Stages of social learning
- Imitation- copy the behaviour of the role model
- emulation- understanding of the goals of action (observational learning)
- facilitation- increased level of effort as a result of others‘ presence
- teaching- Active modification of actions by the role model to improve performance
Outline the link between social learning and tradition
innovation becomes cultural (tradition) when it gets fixed within a population
- innovation is copied by observation (social learning)- then transmission occurs, then becomes tradition when fixed
Outline two types of transmission in social learning
- horizontal- within same generation
- vertical- between generations
Outline examples of foxed vs unfixed innovation (social learning)
Leca et al, 2010- Japanese macaques:
- unfixed- dental flossing- spontaneous innovation, variable techniques, one innovator- didn’t spread in group or to next generations-
- fixed- sweet potato washing, wheat placer mining
- potential reasons for being unfixed- group may be too big, not enough kin, low-ranking female, may not be functional
Link culture and ecological need/nature
- culture is natural process
- exists as soon as opportunity, need and skills present
- exists in many animals
- primate technologies- roots of technological evolution model
Outline human culture
- cumulative culture- gradually improve techniques
- ratchet effect- build on what predecessors have invented, innovations jump from one mind to the next, throughout the population
- potentially more active teaching
- role of written and spoken language
- strong power of normativity
- cultural an biological evolution (dual inheritance)
Define cognition
the mental action or process of acquiring knowledge and understanding through thought, experience, and the senses- involves perception, judgement and action/response
Define intelligence
Ability to easily learn or understand and to deal with new and difficult situations- involves the notions of adaptability and flexibility
Outline cognitive abilities
learning, understanding causality, reasoning, memorising and processing memories, communicating- influences behavioural flexibility, behavioural complexity, modification of actions and environment, manipulation
Outline types of cognition
- Physical cognition- how information about the physical world is acquired, processed and used- involves spatial, timing, and numerical cognition
- social cognition- how information in and from social contexts are acquired, processed and used- involves social learning, theory of mind, and social emotion
Outline methods of studying cognition
- comparative cognition- can compare non-human primates and humans- mechanisms and origins- proximate (process/use) and ultimate (evolutionary relation)
- lab experiments- good as controlled, can specify individual task, can use replicable paradigms; BUT lack of ecological relevance and uses captive populations
- field studies- good as ecologically relevant; BUT observations often anecdotal, hard to control all parameters, ethical issues, experiments hard to realise
Outline the link between brain and cognition
- Primates have larger brains than what would be expected relative to their body size (Roth & Dicke, 2005)
- High energetic requirement- 2% of body weight, but consume 20% of energy
- Expensive to maintain large brains
- large brains associated with higher global cognitive abilities (Deaner et al, 2007)
List different hypotheses regarding evolution of intelligence
- ecological intelligence hypothesis
- social intelligence hypothesis
- Machiavellian intelligence hypothesis
- Cultural intelligence hypothesis
Compare selection pressure na challenges in ecological vs social intelligence hypotheses of cognition evolution
- ecological- ecological election pressure (resources)- challenges are finding, locating, extracting, obtaining resources
- social- social selection pressure (conspecifics)
Outline elements of the ecological intelligence hypothesis
Intelligence/cognition effects:
- where search for food
- where and when memorise food locations
- and how access food times
- involves cognitive abilities of spatial memory, associative learning, and explorativeness
Outline findings on spatial memory as part of the ecological intelligence hypothesis
- Garber, 1989- marmoset use of euclidian map- can remember location of food resources up to 3 years after accessing- mental map can be manipulated and oriented- can travel from any place (not just one direction)
- Normand & Boesch, 2009- similar in chimpanzees (euclidian map)
- Erhart & Overdorff, 2008- prosimians- route-based maps, use of landmark- travel in straight lines, use backtracking, influence of topography- limited in ways to approach but can still manipulate in ones
- De Guinea et al, 2021-black howler monkeys- influence of topography/substrates- Euclidian maps as they arrive at same trees from multiple directions, but contained movement patterns due to limited access of aboreal routes between trees- multiple directions but limited number
Outline findings on searching for good as part of the ecological intelligence hypothesis
- difficulty of cognitive challenge varies based on food sources- e.g. fruits dispersed and seasonal, so greater challenge than leave as evenly distributed and not seasonal
- Harvey et al, 1980- Frugivore species have larger brains than folivore species
- DeCasien et al, 2017- Primate brain size predicted by diet: frugivore > omnivore > folivore
- Malsburg et al, 2020- Habitat generalists (hose that can thrive in many types of habitats e.g. macaques) outperform habitat specialists in cognitive tasks
Outline findings on the extractive-foraging hypothesis as part of the ecological intelligence hypothesis
- Gibson, 1986- Technological specializations in foraging as a result of cognitive abilities of great apes- adaptations for exploiting a variety of high-energy embedded foods through intelligent tool use and its social transmission e.g. dipping, fishing, excavating, opening resources e.g. termites, ants, honey, nut cracking- human evolution as result of year round dependency on tools
- Parker, 2015- feeding requires complexity in processing good- social transmission of these makes up socio-ecological hypothesis- technological dexterity and prowess requires advanced cognition, in new/old world monkeys and great apes
Outline social origins of intelligence according to the social intelligence hypothesis
Dunbar, 1992:
- Individualized relationships organized in a network
- Various relationship attributes- dominance, bond, kinship etc
- complex social networks- macaques (Pasquaretta et al, 2014) and in humans
- Keep track of own and others’ interactions- characterise identity of each individual
- advanced cognition needed for cooperation, competition and social strategizing
Define social intelligence (social intellegene hypothesis)
understanding of identity, kinship, relative rank; showing flexible behaviour according to who does what
List types of relationships explained by social intelligence hypothesis
- dominance
- coalitions
- triadic interactions
- fission-fusion
Explain dominance relationships as part of the social intelligence hypothesis
Bergman et al, 2003:
- dominance relationships and relative rank
- Baboons recognize that a dominance hierarchy can be sub-divided into family groups based on individual attributes and membership in higher order groups
Explain coalitions as part of the social intelligence hypothesis
- more complex decisions based on differentiated relationships due to number of coalition partners as well as observable cues (e.g. large body size)
- grooming reinforces social bonds needed for coalitions (need to monitor own/others relationships)
Explain triadic interactions as part of the social intelligence hypothesis
- Aureli et al, 1992- re-directed agression toward a low ranking kin of opponent (Macaques)- shows understanding of kin relations of other s
- Wittig et al, 2007- 3rd party affiliation- Baboons reconciled on behalf of kin0 increased proximity post-conflict after learning of coalition
- Silk, 1999- macaques- recruitment of 3rd party coalition partners
Explain fission-fusion relationships as part of the social intelligence hypothesis
- Amici et al, 2013:
- variable subgrouping introduces uncertainty- hard to keep track of who hangs out/does what with who
- promotes inhibitory control (separation of intuitions and long term desires)
- F-F species show higher IC
Outline findings on the social brain hypothesis as part of social intelligence hypothesis
Dunbar, 1998:
- Neocortex size (seat of cognitive process e.g. reasoning/consciousness) highly correlated with group size
- not significant relationship with tool extraction (against ecological hypothesis)
Kudo & Dunbar, 2001:
- Neocortex larger in species with larger grooming cliques
- Larger grooming networks in larger groups, larger grooming cliques in larger grooming networks
- suggest grooming clique/network as proxy for social complexity
Deaner et al, 2007:
- Cognitive abilities better predicted by overall brain size than by neocortex size
Als influence of neuronal circuits and pathways, and number of connections
Outline the expensive brain hypothesis
Deaner et al, 2003:
- large brains associated with slow life history
- Life history constrain: slow development and low mortality
- brains: expensive tissue to develop and maintain
- Brain size and life span correlated
Outline the machiavellian intelligence hypothesis
Whiten & Byrde, 1997:
- social intelligence hypothesis is not sufficient to explain higher abilities- social competition as well as interaction leads to machiavellian intelligence
- intense social competition means use machiavellian social strategies to achieve higher social status and fitness
- use of manipulation and deception requires ToM and self awareness - mental representation of others
Outline findings on mental state representation in primates (machiavellian intelligence hypothesis)
- Heyes, 1998- self representation (ToM)- mirror experiemnets- apes yes, monkeys no
- Goossens et al, 2008- others knowledge- Macaques follow gaze of others (understand what others can see)
- Hare et al, 2006- chimpanzee deception- conceal things from others and use hidden routes to access contested food
- Osvath & Karvonen,2012- chimpanzees exhibit self representation, awareness of others knowledge, and false beliefs
Outline the cultural intelligence hypothesis
Tomasello, 1999; Herrmann et al, 2007; van Schaik & Burkart, 2011:
- Social learning is more efficient than individual exploration
- Learning of skills faster with social learning
- Individual repertoire of learned skills boosted by frequent social learning opportunities
- Species with frequent opportunities for social learning can afford to increase cognitive power- means more heavily cultural species should be more intelligent
Outline reserach supporting the cultural intelligence hypothesis
Foss et al, 2016
- zoo individuals (homogenous ecological conditions)- sumatran vs bornean orang-utans
- sumatran more social- exhibited superior innate problem solving, higher inhibition, and more cautious exploration style
- suggests stronger selection on cognitive mechanisms underlying learning in the more social species
Outline integration of hypotheses of cognition/intelligence
- Most likely a combination of ecological and social selective pressures
- Evidence affected by choice of species, variables, analyses and definitions
- Parker, 2015- Extractive foraging hypothesis finds support, but social learning, innovation and tool use also correspond to the cultural intelligence hypothesis
Outline findings on intentionality in chimpanzees
Roberts et al, 2014:
- Language-trained chimpanzees
- Use intentional gestures to coordinate with an experimentally naive human to retrieve hidden food
