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115 Cards in this Set

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V.C. Wynne-Edwards
(1906 -1997)
- Animal Dispersion in Relation to Social Behavior
(1962)
- animals “self-regulate” their populations through
social behaviors.
- for benefit of species
- case for “group selection”, the view that animals sacrifice personal survival and fertility to control
population growth, i.e., for the good of the group as
a whole
- for Wynne-Edwards, animal behavior such as territoriality, dominance
hierarchies, and grouping in large flocks (epideictic behavior) were devices for
the control of population size.
Levels of selection
A. individual
B. kin
C. group selection
Why group selection (usually) won’t work ... (Williams: Adaptation and Natural Selection, 1966)
-By generation 14, over 99% of the population will be producing 3 offspring.
David Lack
(1910-1973)
- 30+ year study of Parus major (great tit)
– reproductive patterns
– studied the factors controlling numbers
in natural populations
Clutch size in great tits
-Size vs. number...
-Average weight of nestlings decreases with clutch size
-Survival of offspring is
related to weight at fledging
Experimental increase in clutch size
-Adult survival decreases with clutch size
J.B.S. Haldane
(1892- 1964)
-While in a pub, Haldane was once asked if he would risk death by drowning to save his own brother. ‘No,’ replied Haldane, ‘but I would to
save two brothers or eight cousins.’
- Wright’s coefficient of relatedness, “r”
r = probability that any randomly selected allele in
a focal individual has copy (identical by descent) in the related individual.
W.D. Hamilton (1936-2000)
– 1964 Journal Theoretical Biology paper
– Kin selection:
- The selection of genes due to an individual’s promoting the survival and reproduction of relatives (other than offspring?)
who possess the same genes through common descent.
– Inclusive fitness:
- The sum of an individuals own fitness** plus all its influence on fitness of relatives other than direct descendants (i.e., offspring).

** Fitness: the contribution to the next generation of one genotype
relative to the contributions of other genotypes.
Hamilton’s kin selection rule
rB – C > 0

r = coefficient of relatedness of donor to recipient
B = benefit gained by recipient
C = cost to donor
Altruism between relatives ...
is kin selection responsible?
- honeybees
– reproductive castes
– defending hive by workers (female)
- monkeys
– agonistic aiding
- ground squirrels & prairie dogs
– alarm calls
– burrow defense
- jackals, red-cockaded woodpeckers, scrub jays
– “helpers at nest”
Kin selection in honeybees?
– reproductive castes
– defending hive by workers (female)

Sex determination in hymenoptera: males are haploid, females are diploid: “haplodiploidy” - more genes in common if only queen reproduces (75% with sisters and 50% with offspring) - queen mates with more than one males, so some sisters could share no paternal genes (queen altered situation - workers locked into choice even if benefit lost).
Agonistic aiding in monkeys
-what is it?
-is it altruistic?
-occurs mostly among kin: is this sufficient evidence for kin selection?
-selfish in fitness enhancing sense.
-difficult to use Hamilton's equation because no precise numbers
-in order to demonstrate kin selection you need to use Hamilton's rule - just association
Alarm calling in black-tailed prairie dogs
A. without close gen rels in home cot.
B. with only non-descendant close gen rels in home cot.
C. with offspring in home cot.

B>A
C>A
B~C

-alerts others but also gives away "alarmers" position - viewed as altruistic perhaps to aid relatives
Cooperative breeders: White-fronted bee-eaters
-average nest size goes up as group size increases
-assist parents in rearing of younger siblings
-manipulate bees to eject poison
Cooperative breeders: Silver-backed jackals
average nest size goes up as group size increases
Cooperative breeders: Red-cockaded woodpeckers
as group size increases so does brood
Florida scrub jay
Helpers at the nest
Woolfenden (20+ yr study)
- defend nest against predators
- defend territory
- help feed nestlings
– provide up to 30% of food
- does not directly increase chick survival: total amount of food is constant; however, it does lessen the burden on
parents.
- helpers tend to be young from previous breeding seasons.
So, why do Florida scrub jays help?
– increase chances of their survival
– likelihood of successful dispersal to a breeding
site is low
– males may inherit all or part of their father’s
territory
– helping increases survival of helper’s parents
– increases the production of siblings
– may result in an increase in territory size
Making the best of a “bad situation.”
White-winged choughs (birds) … in Australia
-apparent altruism that is self-serving
-win over unrelated young
-more individuals in group = can raise more young
Kin recognition mechanisms: Location
– Behavior varies relative to some reference point,
e.g., a nest or burrow (e.g., nestling birds, especially
altricial species). - anyone in nest is treated as kin - opens door for manipulation of brood parasites [error]
- makes room for errors
Kin recognition mechanisms: Association
– Occurs when relatives predictably interact in unambiguous social
contexts (can be based on specific cues like odor; imprinting is one form).
-konrad lorenz and geese [error]
-opens door for errors
Kin recognition mechanisms: Phenotype matching
– Involves comparing the expression of some genetically
influenced trait in another animal with its expression in the individual or others recognized through association.
-Thus it involves learning and the assessment of
phenotypes
-(e.g., sweat bees match the odor of bees entering the
hive with that of bees they were reared with.)
Olfactory cues plus
visual cues...
-Painted wasps' faces and abdomens, altering their yellow markings. Back in the colony, these painted wasps were the victims of considerable aggression.
-- first study showing that wasps can visually recognize kin through facial and abdominal markings and will visually reject unfamiliar wasps.
-Elizabeth Tibbetts, "Visual signals of individual identity in wasp Polistes fuscatus“ (Proceedings of the Royal Society of London B: 269, 2002)
Sherman & Holmes: ground squirrel kin recognition experiment
- Brought to lab at birth and reared under different
conditions:
– siblings reared together
– siblings reared apart
– non-siblings reared together
– non-siblings reared apart
-Later tested in the laboratory as adults
-in the field: full sibs fought less than half-sibs - grew up together
-full sibs more likely to assist one another than half-sibs
[association and phenotype matching play role]
Altruism between unrelated individuals
-Trivers (1971) “reciprocal altruism”
-we help people other than kin, how about animals?
-natural selection ok with favors as long as favor is eventually returned
-cleaner fish - gets food and is not eaten
big fish gets cleaned = not a within species example hard to consider reciprocal altruism because not within species.
Examples in literature:
- Anubis baboons
- Black hamlet fish
- Vampire bats
- Alliances in vervet monkeys
Prisoner’s dilemma and reciprocal altruism
Axelrod & Hamilton (1981)
Conditions:
T > R >P > S and R > (S+T)/2
-disparity
-when would be beneficial for A to cooperate = never
-when only one opportunity to cooperate of defect with some individual
-best strategy = tit for tat - do unto others as you would have them do unto you
-must be able to recognize
Aggression
-A physical act or threat of action by one individual that reduces the freedom or genetic
fitness of another. (E.O. Wilson, 1975)
Agonistic
-Refers to any activity related to fighting, whether aggression, conciliation, submission or retreat.
Aggressive behavior
- “Self-restraint” in
aggression was widely
held to be a trait that
evolved for the good of
the species.
- e.g., Lorenz: On
Aggression (1966).
-vulnerable to selfish mutant
-digger wasps do not use stingers on eachother when fighting for food - Lorenz would say its an evolved trait - not worth dying for
-snakes try to slam eachother to ground - do not use venom - get head to ground
-individual benefits from fighting like this.
What regulates aggression?
- What determines the level of aggression and apparent self-restraint?
- If the object of an aggressive interaction is
to win with maximum fitness gain, the best way to achieve this is to minimize the costs to the victor and not to maximize the costs to the loser.
- Continued attack when it is not necessary to gain an incentive adds risk (injury, etc.).
Aggressive Behavior: Costs
- Potential Injury
- Time & Energy
Aggressive Behavior: Benefits
- Specific benefits are contextually dependent
- General benefits
- Increase individual fitness through access to resources
and/or mates
- Improved reproductive outcomes
- Also reduction of competitor’s fitness
Aggressive Behavior: types
- Territorial aggression
- Dominance behavior
- Sexual aggression (intersexual)
- Infanticide (unrelated target)
- Parental aggression
- Sibling aggression
- Predatory/Anti-predator aggression
Territorial aggression:
Surgeon fish
-Chases rivals from its 2.5 m2 territory 1900 times per day!
Territorial aggression: costly - Yarrow’s spiny lizard
-implant animals with testosterone - look at activity and survival
-with testosterone - active more than others
-1-supplement - testosterone and additional food supplement
2-control = control
3-no supplement - testosterone and no additional food
- testosterone creates extra energy expenditure extra food replenishes
Territorial aggression: Black-winged damselfly
-thought thorax weight created dominant heirarchy - amount of energy - not determinant
-fat content - higher means win (some exceptions)
Speckled wood butterfly – who wins contests?
-intruder loses
-cost to intruder not that great
-concluded got territory=win
-want areas with access to sunlight - sunlight gives them more energy and warmth - intruder does not have this so territorial holder mostly victorious.
Dominance hierarchies
- Schjelderup-Ebbe (~1935)
- Studied aggressive interactions of chickens
- “Pecking order” reflects dominance hierarchy found in groups of chickens
- Unfamiliar hens + fighting over food.
- Usual winner -> “dominant”
- Usual loser -> “subordinate”
- A > B > C > D etc.
- Dominance hierarchies are usually “linear” (i.e., few intransitive relationships) and stable over periods of time.
Dominance hierarchies in primates
- Controversies … how to measure dominance:
-yawning displays?
-grooming?
-access to food?
-access to water?
-access to mates?
-displacement?
-aggressive encounters?
-Correlates of dominance
-infant vs large male - if infant gets food cannot assume the male is subordinate - tolerant?
-"grin" facial expression = submission
Sexual aggression: gulls
-mistaken identity?
-male gets territory for mating - but attacks both males and females - why?
-says male has extra action specific energy territoriality even when inappropriate
-now most disagree strongly
-if not truely ready to mate - she will not put up with attack - taking advantage to courtship feeding
[inter]Sexual aggression: ring dove
-males and females take care of young - even in feeding
-female requires male courtship to come into reproductive condition
-Sperm remains potent in the female’s reproductive tract for about six days.
-when paired with aggressive male female ovulates later than when put with passive male.
-selection shaped males to recognize female condition
-aggression under that condition beneficial to male to verify offspring not their own - previous sperm no longer viable
Infanticide
- Unrelated victims
- Lions
- Langurs; other primates
- Belding’s ground squirrel; other rodents
-killing offspring of competitor - fitness damaged
Austad – Bowl and doily spiders
- Contests for females
- Male lives only three days
- 1st male priority in sperm competition
- 95% fertilized by 1st male
- “Take-overs” do not affect previous mating efforts
- Data show that the average number of eggs fertilized for all females encountered by a male is 10
- Therefore the average value of a female when first
encountered is 10.
- But virgin is more valuable, worth on average 40
eggs
- After 7 min. 90% of eggs have been fertilized so
her value at that time is about 4 eggs.
- Male fighting ability varies with their body size
- Results
- When V (female value) was the same for both males,
fights were settled by differences in male body size.
Larger males won 82% of these fights.
- When males of the same size fought, fights were settled by differences in V.
- When residents were smaller than intruders they persisted more in fights when V was great (e.g., at the end of P.I. period); they gave up, however, after 7 min. copulation. 90% vs. 30% fights resulted in injury).
- Most serious fights occurred where v/k was identical for both contestants (where k=cost of fight).
Game Theory Models
- Models of behavior that take into account that the
benefits (and costs) of a behavior can depend upon what
other animals in the population are doing. They also recognize that, under certain conditions, different
behaviors can be of equal value.
- John Maynard Smith.
- Hawk-Dove model of aggression
- “aggressive” vs. “shy” strategies
Is the ESS (Evolutionarily Stable Strategy) “optimal”?
- No - - -
- Average pay-off is 6.25 per contest.
- If all individuals “agreed” to fight as “doves” (shy) the average pay-off would be 15!!
- But a population composed only of doves would be vulnerable to invasion by a mutant (“hawk,” aggressive strategy).
Brockman & Dawkins
- Female digger wasp
nesting strategies
- “diggers” and “enterers”
- 100 hours to dig burrow
- Females do not seem to
distinguish empty and
occupied burrows – fights
-polymorphism or facultative
use of strategy?
-Measured success of two strategies in terms of # eggs laid per unit time: 68 females; 410 burrows.
-Individual females used both strategies: thus facultative use.
-59% diggers
-0.96 eggs/100 hours
-41% enterers
-0.84 eggs/100 hours
Why sexual reproduction?: Costs
-Costs of sexual reproduction:
- loss of 50% of genes
- energetic cost of producing gametes
- time and energy invested in courtship and mating
Why sexual reproduction?: benefits
-Benefits of sexual reproduction:
- offspring are more varied
- mutations are reduced
Why two sexes?
-Sexual reproduction does not require it: isogamy in
microorganisms; anisogamy in most plants & animals
Sex Ratio
- Sex ratio is usually 1:1 …. why?
- R. A. Fisher (1930) proposed that the common 1:1 sex ratio can be explained in
terms of selection acting on the individual
1:1 sex ratio
-Lots of females in population: advantage to produce males
-Lots of males in population: advantage to produce females
1:1 sex ratio ... exceptions
Acarophenox:
viviparous mite
-Female produces a brood of 1 son and about 20 daughters.
-The son mates with his sisters inside the mother and dies
before he is born!
Mating systems
• Evolutionary view
– Energetics & Economics
• Egg vs. Sperm
– Greater cost of reproduction for the female
– Female is the more “selective” sex (sexual
selection)
– “Ideal” mating systems
• Male acquires as many mates as possible and invests little in parental care
• Female selects “fit” male and leaves parental
duties to him, while acquiring resources required to produce more offspring
Bateman effect: Bateman (1948)
- with more mates, the males mean offspring goes up, while females stays the same.
Sex differences in the maximum number of offspring produced
- males can mate more than once at a time.
-female eggs are more valuable.
Mating Systems: Ecological factors influence how animals are distributed in space
– Polygyny
• Small groups of females
– Promiscuity
• Large groups of females; cues to ovulation lacking
– Polyandry
• When male reproductive output is limited by time/energy constraints or female behavior
– Monogamy
• When environmental demands require both male and female input for offspring survival
Polygyny
•Small groups of females , one male
•e.g., hamadryas baboon; red deer
Promiscuity
-Large groups of females
-Japanese macaques
Polyandry
-When male reproductive output is limited by time/energy constraints or female behavior...
e.g., Sea horse; jacana
Monogamy
-When environmental demands require both male and female input for offspring survival
e.g., gibbons, swans...
Sexual selection
• Selection for traits that are attractive to mates or traits that concern competition for mates.
• A “subdivision” of Darwinian selection that concerns mating.
• Inter-sexual selection
• Intra-sexual selection
M. Andersson (1982): Long-tailed widow bird
- mates increase with length of tail and if tail shortened they lose mates.
Bower birds: Borgia (1982)
-build open ways through grass as attraction for female.
-Males with more decorations get more females
Sexual Selection Hypotheses
• R.A. Fisher: “Runaway” selection - most broad and applicable
-randomly occuring mutations for female preference - mutations in males to accomodate females - in some instances the trait is linked with fitness trait - once you get to particular point where large enough - group of female will prefer one trait - if you don't have it possibility of mating decreases
• W.D. Hamilton & M. Zuk: Disease/parasite avoidance (narrow version)
-stickleback fish - bright colors prefered - fitness enhancing component (disease/parasite avoidance) - must mean good health - smaller subset of characteristics
• A. Zahavi: Handicap principle
-i.e. longer tails means better flight - genetics moves toward that - female preference genes creates pressure to evolve - want survival advantage and female preference - optimal length - traits that evolve and are preferred by females must be costly to male - if traits were easy they could just produce it - but if costly then only truely fit males can have it - says "handicaped" because so costly male must be very fit - males not as fit cannot bear burden and pull it off.
Results of intrasexual selection on males: Adaptations that enable males to compete for reproductive success
• Guarding behavior
• Sperm plugs
• Biochemical substances
that reduce female receptivity
• Low threshold for mating
attempts
• Monopolization of females
• Postponement of mating
attempts
• “Sneak” copulations
• Forced copulations
• Sexual interference
• Female mimicry by males
• Assaults on male competitors
• Assaults on offspring of
competitors
• Size of testes
Damselfly mate guarding
add notes.
Love bugs
-56 hour copulation tie
Orb-weaving spider ( Argiope aurantia )
The male of this orb-weaving spider self-destructs during mating. By dying while still joined to a female, his corpse forms a kind of “chastity belt.”
The dead male's inflated palp
remains wedged inside the female. This prevents her from mating again, and gives his sperm time to fertilize her eggs. The female cannot pull the male out for at least 15-25 minutes. Other males try to pull the dead male out, but most often fail.
(Like many spiders, the female eventually eats her deceased mate.)
Biochemical warfare: Heliconius erato - butterfly
-During copulation, male deposits scent that renders female unattractive to other
males.
R. Thornhill: forced copulation
in scorpionflies

Forced copulation also in water fowl (ducks), and orangutan
- male will not accept rejection from female.
Female mimicry by males
Desmond Morris (1952)... “homosexual behavior in the three-spined stickleback”
Assaults on male competitors
-Assaults on offspring of competitors
Relative testis size in primates
-Circle size = relative degree of sexual dimorphism
-Shaded oval = relative size of testes
-Arrow = relative size of erect penis
Parental care
- Correlated with increased
parental care is a reduction
in the number of offspring:
- “r” selection: many
offspring, little investment
- “K” selection: fewer
offspring, greater
investment
Parental care in the stink bug
-About 28 eggs produced.
-Female remains with eggs over two weeks.
-Protection against predators/parasites.
-Remove female and NO eggs survive; with protection 50% survive.
Parental Care brief survey: Teleost fish (bony fishes)
- 79% No Parental Care
- In the rest, usually one parent involved
- Guarding the eggs
- Fanning the eggs
- Internal fertilization: 86% Female Care
- External fertilization: 70% Male Care
- Paternal certainty?
- Order gametes released: who gets to leave first?
Colony of breeding male bluegill sunfish: Neff, B. D. Nature 422, 716-719 (2003)
-Parental males are tending their nests, while females ready to spawn are swimming higher up in the water.
-Neff showed that male bluegills use two cues to assess how likely it is that they are the father of the offspring: the more certain they are of their paternity, the more attentive they are as parents. They are also more likely to defend the nests against an egg predator.
-Two distinct male reproductive strategies have evolved. Males termed 'parentals' defend nest sites, attract females, and then care for the eggs and newly hatched offspring. Other males mature at an earlier age as 'cuckolders' and steal fertilizations from parentals either by darting into nests at the critical moment of spawning ('sneakers') or by mimicking females, apparently fooling the parental male into ‘thinking’ he has attracted two females at once. Sneakers
are particularly effective, fertilizing 89% of the eggs released by a female during the 8% of spawnings in which they participate.
-The presence of sneaking thus provides a cue that parental males could use as a guide to their paternity. The second cue is more surprising, but has been confirmed by controlled experiments: parental male sunfish can apparently assess their relatedness to newly hatched fry using water-borne odor cues; the mechanism is
unknown, but other studies of fish suggest a role for genes in the major histocompatibility complex (MHC) in olfactory discrimination of kin and
potential mates.
Colony of breeding male bluegill sunfish experiment
In the first experiment, parental males that were spawning were exposed
visually to four sneaker males, enclosed in transparent plastic containers so
that they could not fertilize any eggs. Control males were exposed to empty
containers. As predicted, males reduced their level of care during the egg
phase when they were tricked into expecting lower paternity.
Then, male care was tested a second time after the eggs hatched, when the
second mechanism for assessing paternity —olfaction — was predicted to
restore the certainty of paternity of the experimental males. This is exactly
what happened: these males increased their care when the new information
suggested that their paternity was not lower than in the control group.
These reductions and increases in care fit perfectly with predictions based on
the two mechanisms of assessing parentage.
Reliability of paternity and male care
- Hypothetical case
- If % of eggs fertilized by male when he copulates is
low (say 0.4) will paternal care evolve?
- Perhaps, if:
- Survival of offspring produced offsets the loss of
additional females
Factors Promoting the Evolution of Parental Care
- Stable, complex environments
- Difficult environments (major & extreme fluctuations)
- Specialized Diets
- Predator pressure / infanticide
Parental Investment
- As defined by Trivers (1972)
- Any investment by the parent in an individual
offspring that increases the offspring’s chances of survival (and hence reproductive success) at the cost of parent’s ability to invest in other (future)
offspring.
Reproductive Value
- As defined by R.A. Fisher
- Number of offspring remaining to be born to the
average female at age ‘X.’ Reproductive value changes with age.
Maternal investment (MI) in wild horses: effects of female reproductive value? Cameron et al. (2000)
Older mares were more protective for the first 20
days of life, but less diligent thereafter. Total MI
by older mothers did not seem to be any greater, but was better targeted at the most critical period for foal survival. Older mothers were also more likely to foal in consecutive years, supporting the hypothesis that they are investing less than younger mares in individual offspring.
Communication: definition
action* or feature of one organism that alters the probability of the behavior of another organism in a fashion adaptive to either one or both
of the participants. (E.O. Wilson, 1975)
* can be the product of natural selection and /or learning
Emphasized modality and phylogeny
-no relationship
-chemical, visual, auditory
Signal modality
variable benefits & costs
Habitat influences on great tit song (Hunter & Krebs, 1979)
Forest: narrower range
of frequencies; lower max.
frequency; fewer notes
Bird alarm calls
-alarm calls of birds are all found in the same range so that all may benefit from the knowledge of an enemy.
Discrete vs. graded signals
-[zebra ears] - there is happy and angry and then degrees of that.
Ritualization: the origins of signals - definition
the evolutionary modification of a behavior pattern that turns it into a signal used in communication (or improves its
efficiency as a signal)
Comparative evidence for the ritualization of a display
a. male domestic fowl
b. ring-necked pheasant
c. impeyan pheasant
d. peacock pheasant
e. peacock
Communication: information sharing or manipulation?
Dawkins & Krebs (1978). Animal signals: information or
manipulation?
“One party - the actor - emits a signal, to which the other party - the reactor - responds in a way that the welfare of the species is promoted.” (Tinbergen, 1964)
“Displays are acts specialized to make information vailable.” (W. J. Smith, 1977) The Behavior of Communicating
Batesian mimicry
harmless species mimics harmful one Müllerian mimicry: harmful
species look alike
Discrete vs. graded signals
-[zebra ears] - there is happy and angry and then degrees of that.
Ritualization: the origins of signals - definition
the evolutionary modification of a behavior pattern that turns it into a signal used in communication (or improves its
efficiency as a signal)
Comparative evidence for the ritualization of a display
a. male domestic fowl
b. ring-necked pheasant
c. impeyan pheasant
d. peacock pheasant
e. peacock
Communication: information sharing or manipulation?
Dawkins & Krebs (1978). Animal signals: information or
manipulation?
“One party - the actor - emits a signal, to which the other party - the reactor - responds in a way that the welfare of the species is promoted.” (Tinbergen, 1964)
“Displays are acts specialized to make information vailable.” (W. J. Smith, 1977) The Behavior of Communicating
Batesian mimicry
harmless species mimics harmful one
Müllerian mimicry
harmful species look alike
Eggs of cuckoos and their hosts
- cuckoos try to match eggs outer appearance exactly to hosts.
Deception: “femmes fatales” fireflies
-Female Photuris firefly devours a male Photinus ignitus
-female learns lighting patterns of females of other species and mimics them and kills males she attracts.
Plover broken-wing display
-deceive predator and lead him away from grounded nest by pretending to have a broken wing.
Beau Geste hypothesis: Mockingbird song
-beleived to mimic other birds to keep them out of territory by making them think the area have too many birds in the location already.
Sentinel birds (C. Munn, 1986): antshrike and shrike-tanager
add notes.
Monkey alarm calls: Vervet monkeys and Japanese macaques
- the lower "class" monkeys fake monkey calls while dominant are feeding so the other run away and the monkey can feed and not wait.
Selection may promote signals that are highly reliable
and cannot be ‘faked’
- frog size coordinates with pitch of vocalization
Harris’ sparrow (Rohwer, 1978)
-having a large amount of colored breast and head feathers puts you in higher rank and rank decreases with amount of color
Stotting
- gazelles use this to try and show off their fitness to predators to tell them they are too fit to eat.
-will not do this with cheetahs because cheetahs are sprinters and there would be no time to display. - will do with dogs (more long distance)
Skeptical response in rhesus monkeys (Gouzoules, Gouzoules & Miller, 1996)
-Rhesus monkeys habituate faster to the false alarm calls of lower-ranking group members (still habituation with high ranking members, but just occurs faster with lower ranking monkeys)
Vervet monkey alarm calls
-vervet monkeys have different alarm calls for different predators.
-leopard - climb tree and go to terminal branches where leopard cannot reach - do not give alarm call to any other cat/canine but leopard.
-eagle - retreat to inner branches where eagle cannot reach without damaging itself
-heard calls and gets representation of threat and react accordingly
-young monkeys must learn differences between random species and predator -elders do not respond to younger vervet calls without checking first.
-snakes - not concerned by cobras, but by pythons
-learning component to calls
D. Griffin (1915-2003)
- 1976
- “The Question of Animal
Awareness”
- 1984
- “Animal Thinking”
- 1992
- “Animal Minds ”
- 1982
- “Chimpanzee Politics” F. de Waal
(Note: ape “language” studies of 1960s
predate the revival of a more general
interest in animal cognition.)
“Animal Cognition”
-emphasis on flexibility
-facets of intelligence
-Learning
-Remembering
-Problem solving
-Rule and concept formation
-Perception
-Recognition
“Clever Hans”
- Psychologist Oskar Pfungst
showed Hans was picking up subtle unconscious cues
from people asking questions.
Categorization in Pigeons
- R. Herrnstein (early 1960’s)
- Showed pigeons in lab slides of outdoor scenes, reinforced pecks that were given if the scene contained a tree.
- The scenes showed many different things in addition to trees (cars, people, buildings, etc.).
- Tested birds with large number of slides after learning phase
- They could remember scenes with trees they had seen after months!
- They could accurately (75% correct) classify a large number of new slides.
- The pigeons had apparently created a rough mental category for trees.
- The “mistakes” often turned out to actually have trees in them (distant etc.) or sometimes had telephone poles in them!!
Do monkey calls have meaning to the animals themselves?
Cheney and Seyforth: habituation and dishabituation technique
-present stimulus to non verbal baby - habituation arises
-present another stimulus - will it classify stimulus as 1st? or a more vigorous response? - treats it as something different
-are the wrrs and chutters of when other groups approach or become aggressive the same response or different - like eagle and leopard alarm calls
-the wrrs and chutters have same meaning - habituation
-the eagle and leopard have different responses and are not habituated.