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what impacts seasonal fluctuation
seasonality is dependent on rainfall, not heat
reduction in food intake - orang 85-90%, shifakas - 74%, gorilla - 20%, chimp - 46%
how do they deal with seasonal fluctuation
switch food, typically to lower quality
shifts in ranging, ranging in unknown areas —> increased energy expense (traveling farther), less efficient foraging (dont know area)
changes in grouping patterns, especially fission-fusion
dietary shifts are most common because the others have very high risks, but larger bodied primates are more likely to shift ranging patterns, lactating chimps seek shelter in caves to cool down
fallback food
anything consumed when the main food is not available
negatively correlated with abundance of preferred foods
allows for reproduction even in unfavorable conditions
reproductive seasonality
reproducing during certain seasons. most wild populations have at least some degree of reproductive seasonality
icome-capital breeding framework
an income breeder requires daily reaching of caloric intake; reproduction varies day-day
capital breeder can rely on fat when food is low, allowing for reproduction even when food is scarce
primates sit in the middle and can change their diet. Middle to late lactation usually falls with the greatest food abundance because it is very energy intensive
when there is a short time where females are fertile, there is no male competition because females cannot be monopolized
sentinel species
tell about the health of the ecosystem
flagship
charismatic megafauna
ecosystem services
species does a job, such as seed dispersal
spatial ecology
home range size and home range overlap
home range size
space the animal will use over a specific period of time; can change seasonally. typically larger with body size and males. group (cumulative) mass, instead of individual, influences range
arboreal species and overlapping ranges have more of a cumulative body mass relationship; frugivores have larger home ranges than folivores
infant carrying
unique to primates except for marsupials; hiding could be effective against predators in most species; large litters provide thermoregulatory benefits and makes them impossible to carry, so they are parked in nests
home range overlap
large level fission-fusion for predator defense; ecological knowledge + sharing of information for enhanced foraging; increased resource holding potential because of temporary larger group; reconnaissance before transfer (learn about another group before joining it; extra-group mating for genetic diversity
territory
portion of a home range actively defended
what influences territoriality
territorial males have a smaller range which is defended for use in mating, feeding, and all activities. It is more limited in primates than other mammals, which could be because it reduces the costly aggressive interactions
maintained through scent marking, patrols, visual rituals
simple optimality model for territoriality
predicts territory size has to be balanced by benefits and costs of defense. only species that can travel far enough ina. day to visit the entirety of their territory should be territorial
chimpanzees are exceptions, they have large group size and fission-fusion which allows them to have a larger defended territory
mate defense hypothesis
monopolize a mate. contest shoul dbe higher when estrous female is present; high female-male ratio, males should her females, aggression should be largely intrasexual
male resource defense
males defend resources for females; dependent on habitat quality, so males are more territorial to monopolize it; more imprtant in single male groups
dispersal meaning and types
emigration from a social gorup
primary - from natal group
secondary - dependent usually on demographic levels
trends in dispersal
in mammals, male biased
in primates, female biased
costs and drivers of dispersal
primary cost: aggression from new group; minimized with parallel dispersal
drivers:
ecological; lower food availability=leave; (chimps actually leave when food is more abundant)
physiological - sex hormones (if leaving at sexual maturation)
social: conspecific attraction (leave during mating season), agressive eviction (remove males from harem), reduced social integration (less grooming, avoidance)
why would you disperse?
inbreeding avoidance
local mate competition - sex that competes more strongly for mating opportunities should disperse
resource competition - pair bonded species - finite resources, only one breeding pair
why would you oppose dispersing?
hyperlocal knowledge of food resources
mortality costs
loss of kin cooperation
why is there female-biased dispersal in primates
dominant males have very long tenures, overlap between dominant father and daughter reaching sexual maturity
males benefit from kin cooperation - hunting and territoriality (ex: chimps)
collective action problem
coming to a consensus as a group
pay something to be in the group, which is a cost to some; giving up selfish perspective
group size paradox
the larger the group the more likely they SHOULD be to make a decision, but in actuality collective action problems are more pronounced because each individual is more likely to defect
territorial defense becomes MORE of a problem as the group gets bigger
there are logical or informational constraints on getting the relevant individuals to act in concert (insufficient incentives to get individuals to participate)
solving collective problems
exploitation of the great by the small: most dominant makes the decisions (strongest/biggest/highest caloric intake = more incentive to move); highest rank leads efforts
dominance asymmetry, coercion, most knowledgable
dominance asymmetry in solving collective problems
individuals will follow others, so only x number of individuals are needed for the whole group to follow
coercion by dominant individuals in collective problems
Unlikely to work in practice because of the inability of a dominant animal to force a consensus decision, either because:
It is physically unable to do so, particularly in large groups
It would not gain sufficiently to outweigh the costs of coercion
collective problems solved by the most knowledgable
mostly in humans than nonhumans, but is seen in groups led by older individuals
consensus decision making
members of a group vote (different calls can mean different things); depends on rank and number of individuals voting.
individuals forego personal interests for group interests (reduced fitness); consensus costs can be so pronounced that they lead to segregation (breaking up of the group based on needs); they will stay together with. hugh predation because they have shared info and incentives
disagreements in initiators for which direction to go leads to anchoring where the group wont go in any direction
what influences who will take leadership
motivation - dependent on resource need; sexually dimorphic males = males have more motivation; females have to reproduce = more motivation; whatever is most energetically demanding = more likely to be leader
temperament - individuals who talk/vocalize more have more leadership potential
knowledge - age correlates with leadership in areas requiring specialized knowledge, but not when risk-taking or physical bravery are needed
dominance status - more resource holding potential
what is a social bond
interactions among individuals, relationship between them, social structure all relate to each other
top grooming partner - grooming exceeds time needed to remove parasites
strongest bonds are between kin; nonkin bonds are called friendships
poking eyes is a trust exercise between capuchins
how do we describe social relationships?
frequency, diversity, symmetry, tenor, tension, predictability, stability
what is the adaptive value of social bonds
in baboons, increases infant survival.
highest bonds are with mothers and daughters, and then sisters. females with more female bonds have higher survival rates
social bonds for males
strong bonds are tied to future dominance rank. dominance rank is tied to paternal success. so, social bonds are tied to paternal success
if bonds are not maintained, then neither is rank or reproductive sucess
social bond research topics
reproductive rate - if strong bonds are important, reproductive rate is higher
Offspring survival - more infants survive
Longevity - lifetime reproductive success or mortality based on long term social bonds
The best predictor of how many offspring is how long they live
social relationships and mortality in humans
close social relationships help survival so much it can offset smoking
social support is good all of the time
access to resources, agonistic interventions, predator avoidance, stress buffering
stress buffering
glucocorticoids (stress hormones) are good in the short term because they bring stored energy into use. but, theyre bad in the long term because they cause decreased immunity, cardiovascular problems, and lower reproductive hormones
social interactions can decrease stress hormones and buffer against stressful situations
social stress and dominance rank
higher ranking individuals have lower stress hormones, especially for high ranking females (less competition, access to food)
alpha male stress is higher depending on the stability of the dominance hierarchy; more unstable for males
intergroup encounters raise stress levels. even in acutely stressful situations, bond partners will reduce stress hormones
proximate mechanisms for social behavior
mate access, social status, stress buffering = higher success
what is cooperation
benefits recipient but can be costly or beneficial to the actor
mutualism
both actor and recipient benefit
altruism
benefit recipient and carry an immediate cost to the actor
what are some constraints of cooperation
cognitive constraints: keep track of costs and returns over minutes vs years
defector coming into a group will always have higher fitness than the cooperative population; due to breeding, overtime the whole group can become defectors and the grouo will fail
cooperative behavior in nonhuman primates
almost all of it is kin selection or direct reciprocity
benefit*degree of relatedness > cost
Who should you bias your behavior for based on relatedness alone?
the one with the highest percentage of your genes, typically your offspring
kin recognition is required for kin selection. how do they recognize?
close association and familiarity
depending on how many males (harem - offspring are all related vs multi male); fewer males (high reproductive skew) results in higher tolerance because it is more likely they are all siblings
phenotypic cues - olfaction in steps, owl monkeys
reciprocal altruism (direct reciprocity)
same individual must assist repeatedly; frequency they give is contingent on frequency they recieve; there are temporary net costs; it relies on payback AT SOME POINT; they cannot be close relatives or prospective mates
cheaters (ones who wont give and only take) must be detectable
sense of fairness
capuchins and chimpanzees are less likely to do work, exchange, or accept a reward if another individual got a better deal
indirect reciprocity
benefits are not gained from the individual helped, but from a third party observing the interaction. good reputation, feel good activity, a good reputation is more valuable than money
network reciprocity
cooperators form clusters where they can earn higher payouts than defectors
group selection
selfishness beats altruism, but altruistic GROUPS beat selfish groups
kin selection
choosing to support kin over all others
what are the types of altruism
kin selection, direct reciprocity indirect reciprocity, group selection, partner choice
all must weed out defectors, or risk the group falling apart
two kinds of sexual selection
males compete for a mate (intra)
females choose a male mate based on appearance (inter)
intersexual competition
mammalian females are usually the choosier sex
choose for: paternal investment, territory, good genes, maturity, dominance, weaponry (fighting ability and rank), male investment, symmetry, vocal displays, olfaction, heterozygosity
testosterone
males with high testosterone are more aggressive and more colorful
direct benefits
territory, provisioning, paternal care
indirect benefits
good genes, MHC diversity
good gene hypothesis
elaborate male traits may be indicators of heritable genetic quality; this relies on honest signals and handicapping traits.
males have a trait that reduces viability (ex: bright coloring), and survival despite this handicap indicates high quality genes, so females will mate with brighter colored males
with trait still have same lifespan and more offspring survive
sexy sons hypothesis
not often used in primates
trait has evolved beyond being an honest signal, decreasing the fitness of the male; females ONLY select so their sons have that trait and can get more mates
fisherian runaway selection; correlation between trait and female preference —> females choose the modified more dramatic signal over the real males
lifespan is shorter and offspring do not survive at a higher rate, but they do have more mating opportunities
compatibility hypothesis
mechaism for good genes: Major histocompatibility complex (MHC); selects for immune function
complementary or very different MHC proteins would enhance and offspring’s immune system, so mates are chosen to diversify immune systems
positive correlation between mating and different MHC; golden snub-nose monkeys are thought to have MHC-biased mate choice because their MHC is more diverse than it should be
what are the three hypotheses for indirect benefits
good genes, sexy sons, compatibility
polyandrous mating
females appear to go out of their way to mate with more than one partner (brown capuchins, ring-tailed lemurs)
pros and cons of polyandrous mating
risks: harassment from dominant males, STDs
pros: lower infanticide risk - if they dont know who the father is, they will think it is their own (paternity confusion)
paternity confusion hypothesis: required that females cannot be monopolized; females can avoid being monopolized by unreliable timing of ovulation and sexual swellings
male mating strategoies
sexual coercion, reproductive concessions, consortships and mate guarding, male infanticide, male-female friendships
why are male-female friendships beneficial
females prevent infanticide and males get future mating opportunities
basic transaction model (concession model)
males can control reproduction but presence of subordinates increases fitness; they allow subordinates opportunities to mate in return for defending the group from loner males trying to overthrow the alpha
reproductive skew models; seasonal breeders?
basic transaction model and compromise model
seasonal breeders have minimal reproductive skew since females cant be monopolized
compromise model
dominant is unable to control reproduction and exclude subordinates; this decreases the groups productivity
male consortship and mate guarding
males that herd or maintain proximity to a female when she is fertile, are more likely to sire offspring
mate guarding is aggressive to other males, and is not female choice because all other choices are prevented. alpha male should choose older females who have high reproductive success
male infanticide
widespread convergent evolution, but most common in social species, and almost absent in pair bonding (callitrichids - one reason monogamy evolved)
occurs more with higher reproductive skew toward the alpha male, because subordinates know its unlikely to be their offspring, and with longer tenure length following takeovers
shortens period of postpartum infertility, allowing males to impregnate more females more quickly
female counterstrategies to male infanticide - evolutionary arms race
promiscuity - no one will know if theyre the father
stay in the proximity of potential defenders
female coalitions - work together to run off males too close to an infant
group splitting - when a takeover happens, those without infants remain and those with leave with the former resident male
Bruce effect
what is the bruce effect
if females are pregnant when a takeover happens, they spontaneously miscarry, which is thought to prevent infanticide
stress hormones increase and estrogen plummets when new male comes into the group
challenge hypothesis
the role of testosterone in male-male competition: musculature, armor, body sexual dimorphism, fear reduction, vigilance towards dominance-related behaviors. however, it is taxing on heart muscles, so it has to be tightly moderates to avoid death
mandrill face color is associated with high testosterone
testosterone should be the highest during mating season for seasonal breeders, or when competing for a fertile female
high rank predicts mating access, so high ranking males should have higher T
T is higher during group instability
sexual dimorphism + social mating groupings
larger dimoprhism = intense male-male competition, polygynous mating systems, low levels of male parental care
less dimorphism = social monogamy, higher levels of male parenting, less male-male competition, lower testosterone
correlates of paternal care
Social monogamy in species with long-term pair bonds\
Territoriality in which groups defend an exclusive area
Reduced sexual dimorphism when the difference between male and female size and traits are less pronounced
parental care
anything that is likely to increase the survival and fitness of young
parental investment
investment by parent that increase an individual offspring survival but at the cost of investing in other offspring (foregoing mating, reduced parental survival)
which offspring should be invested in?
healthier, bigger, younger individuals, and males because of reproductive asymmetry
when should offspring be invested in?
if benefit to parents is greater than the cost
future prospects of reproducing are low
successful rearing requires two parents
decrease investment if relative value of offspring is low, or presence of alloparents offsets costs
females have a lot of investment already at time of birth from carrying the fetus, so they will continue to invest
when are alloparenting and biparental care more common
when there are long periods of offspring dependency
orangutan interbirth interval
8-9 years
life history stages
infant - until weaning
juvenile - until puberty
adolescence - puberty
adult - first reproduction
maximum life span drive reproductive success in females
r selected
many offspring with low investment and low survival (RABBITS)
k-selected
few offspring with high investment and high survival (KANGAROOS)
primates are K selected
Slow life histories, altricial (needy) offspring, single births (except streps), long gestation
how long is elephant gestation
2 years
why is childhood unique to humans
adults still provision for children, while juveniles in primates do not get provisions after weaning
male vs female metabolic expenditures
energy is finite and must be spent on somatic functions (growth + maintenance), survival, and reproduction
males sacrifice growth for energy to spend on play
females play less so they can use more energy for growth, and reach adult body size faster
major adaptive hypotheses for play
motor skills, training for the unexpected, social skill development
why do males play more
motor skills are needed to defend a territory/females and move to the top of the dominance hierarchy
unexpected physical altercations
formation of social bonds - philopatric sex might play more
factors influencing primate life history
mortality rate, infanticide risk (if high, mothers might accelerate weaning), diet (+seasonality), encephalization
natal coats and infanticide
natal coats signal to group members that infants should be treated differently (oddity efect)
cryptic color to avoid predation
warmer for heat retention
infanticide is a driver of neonatal coloration change, as mothers will wean earlier to change the coat color so theyre not targeted
ecological risk aversion hypothesis
assumes that juveniles are less efficient foragers and position themselves near conspecifics to avoid predation, and thus experience high levels of intragroup feeding competition
evidence supports that they occupy central positions, but by weaning they already have adult foraging abilities
needing-to-learn hypothesis
correlation between brain size and reproduction age - essential skills are learned during development and required for adults
opportunities for socialization and long term relationships
why do primates have a protracted juvenile period
social skills development
needing-to-learn
ecological risk aversion