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bio 472 final


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What are the 2 types of ANTIPREDATOR BEHAVIOURS?
Primary Defense:
-avoid detection in the first place

Secondary Defense:
-avoid capture once detected
What are the methods used in PRIMARY DEFENSE?
1) Be randomly placed in habitat to avoid search image formation and area restricted seraching
---eg: Ptarmigan nest placement. Puts it in cover (rocks and trees) to prevent learning of preds, good idea to switch it up randomly.

2) Have a different period of activity than predator or hide in inconspicuous place

3) Crypsis
-look like environment (camoflage)
-color, pattern, disruptive coloration
-behaviour: posture and background matching
-*Costs of Crypsis: i) must be still therefore energy intake lowered; ii) change body shape and behaviour to imitate something you aren't
-don't storytell, animals senses may be different from ours

4) Aposematic (Warning) Colors
-opposite of crypsis. Stands out from environment to signal that you are no good
-How does it evolve? i) advantage to individual(if it escapes); ii)advantage to kin

5) can lead to Mullerian mimicry. ie: diff spp (both posonous evolve the same colors to take advantage of same system

6) Batesian mimicry (model is toxic, mimic is tastey)
What are the methods used in SECONDARY DEFENSE?
1) Withdraw to cover (natural or a den/burrow etc)

2) Rely on speed to escape
-must have enought time -> vigilance
-*vigilance varies with group size
-*vigilance varies with predator densities
-*vigilance varies with position in the group (ones on edge have increased vigilance)

3) Startle response or confusion effect
-erratic escape path
-bright markings or colors help to confuse -> called 'FLASHING'... overwhelms the predators mental capacity
-can be paired with crypsis. look cryptic as long as possible, then flash to escape

4) Diverging
-dispay (eg: broken wing display of Killdeer) to divert attention from eggs
-colors and behaviours to divert attention to less vulnerable body part

5) Bluff or intimidation
-looking big/aggressive
-play dead

6) Active Defense
-teethe, chemicals, claws, horns, etc
Benefit to Individual
-invite pred to pursue (from a safe distance that will fail)
-inform pred that it has been seen, therefore deter puruit
-ambush hypothesis. High leaps allow overview of predators and landscap to prevent ambush
-signalling of health (deter puruit). eg: individuals stotting at a high rate had fewer preds attacking them
-startle effect
-confusion effect (within a group)

Benefits to Group Members (conspecifics)
-increases group cohesion to aid the confusion effect
-mothers signalling to offspring... distracts attention from offspring
-the role of hormones in behaviour. Proximate perspective
-hormones are slower messages than electrical (ie: nervous system)
-there linked to long term control of vehaviour
-transmitted through the bloodstream
What are some ROLES OF HORMONES?
1)Important in Development
-ie: sexual behaviour in rats
>parental care roles, mating behaviour, copulation etc. is linked to different hormones in adults
>if gonads of adults are removed, they don't show typical mating behaviours
>if you inject testosterone or estrogen into such rats, they regain typical behaviour
>if you infect opposite sex hormon into rat, they won't behave like opposite sex
-rat fetuses are sensitive to hormones with respect to brain development 15-27 days post conception. Even small doses may be important
-birds... can the mother adaptively allocate testosterone to eggs?
>ie: last eggs to be incubated (ie: the runts) get the testosterone to make up for the disadvantage)

2)Hormones affect Morphology
-eg: toads, androgens increase number of muschle fibers in larynx, enables the 'trill call' of courting males
-eg: antler size of deer is partly influenced by hormone levels
-eg: birds>selected plumage, if gonads removed, birds will look like males

3)Regulate Day to Day Behaviour
-eg: fight or flight controlled by epinephrine and norepinephrine, made in the adrenals, controlled by the hypothallamus, stimulated by a stressor
-it is difficult to determine cause and effect (ie: does testosterone cause more victories, or does more victories cause more testosterone)
-eg: male squirrel monkeys
>ambient testosterone levels of male prior to grouping with other males DID NOT predict eventual dominance
>after dominance settled, dominance had higher testosterone levels

4)Constrain certain behavioural options
-eg: birds with high T = high territoriality and high mate searching & polygeny
In terms of Hormones, what does stress do?
-impacts levels of corticosteroids (ie: glucose metabolism, immune sys, etc.) Inhibits reproductive bevaviour, decreases steroid hormones
-Testosterone inhibits immune system
-eg: mice
>dominants traded off survival for short term reproductive investment (high T -> defended territory and females)
>subordinates prolong life and get as many chance matings as possible
-hormones that are impacted by stresses

-eg: Kittiwake chicks experiment
-food shortage = increased corticosterone level in chicks = increase in begging and aggression = increase food delivery by parents = decrease in cortical sterols
-if food shortage is chronice, then chicks develope with long term high levels of cort... = long term begging behaviour
What is the best way to study memory /learning?
-general process and adaptive specializations

eg:Clark's nutcracker
-one of the most remarkable food storers
-buries seeds in soil
-can cache between 22,000-33,000 seeds in up to 9,000 locations in one summer, therefore must remember locations

2 types of storage:
i)Larder hording: lots of food hidden in one place
ii)Scatter hoarding: what nutcrackers do

How do they do this?
-random search? no. too slow
-olfactory searth? no. Nutcrackers only find own seeds, this will lead them to others
-caching algorithm? no. they don't depend on constant positioning (ie: south of rocks)
-memory for sites? YES

Nutcrackers use spatial memory via landmarks
-expt: 1/2 landmarks shifted after bird had cached
-birds searched 10cm from landmarks therefore must use objects distant to chache as well (makes sense because think of changes in environment that occur from sumer to winter)

-hippocampus key brain area in learning/memory (lesion studies important
-storing birds have larger hippocampus volume than non storing
What are the 4 ways of studying sensory systems?
1) Directly record neural activity of sensory neurons
-impulse freq associated with different stimuli: "can it detect this stimulus?"

2)Block or interrupt sense organs or neural connections (chemically, surgically, mechanically [eg: blinders in pigeons])

3)Learned Responses
-try to condition an animal to respond to a stimulus (eg: ringing or bell)
-then change volume gradually and see thresholds where there is no response

4)Natural Responses to Environmental Stimuli
-eg: how acute is vision based on how the animal moves to or from objects in the environment
When there is too much happening to conduct a good study, what 2 types of filtering are available to help?
1)Peripheral Filtering
-at level of sense organs
-limited in what range they can respond to
-eg: male tree frogs have a "co-qui" call
>males hear the co, females hear the qui
>this is due to different tuning of eardrums in the sexes

2) Central Filtering
-occurs at the level of he nervous system; brain ignores certain input
Why do Animals have particular senses?
-stimulus properties have different costs and benefits depending on environment
-(ie: you can interperet light faster than chemicals)
-stimulus properties vary with the environment
What is photoreception?
-use of light
-short wavelengths = UV, usually causes damage to organic molecules
-long wavelengths = infra red (thermoreception), insufficient energy to be useful
-UV is common as a signal of quality in plumage of many birds, used in mate choice
-ppl see 400-750nm
What is Infrared reception?
-percieved by relatively few animals
-see in heat, like thermogoggles
-most common in snakes (have pit organs to detect heat)
-strikes at warm prey
How is Chemoreception used?
-needs to identify the chemical (what type is it?)
-needs to distinguish concentration (sense direction, distance, size of organism etc.)
What are the main types of senses used in animals?
What are the 2 types of Semiochemicals?
1)Pheromones (intraspecific)
-releasers; quick acting, affect behaviour
-primers; slow acting on physiology

2)Allelochemicals (interspecific)
-allomones; toxins, lures, repellents etc (benefits the sender)
-kairomones; attractants, alarm signals, etc (benefits reciever and sometimes sender)
Give an example of Human Chemoreception.
-can tell diff of sex based on breath
-kids can diff shirts worn by kin vs stranger
-moms can ID shirts of own kids
-women can synchronize PMS
What are the 4 types of Mechanoreception?

-detects position and motion of body parts. Stress and tendons and ligament

3)Balance and acceleration
-eg: statocysts in invertebrates semicircular canals in ears in vertebrates

-a)amplitude (loudness)
-b)Frequency (pitch)... humans can hear in 20-15,000Hz
-c)temporal patterning of amplitude and frequency
What are the 3 types of hearing?
1) Directionality Hearing
-timing of stimulus contacting 2 sense organs (ears)
-most animals are good at horizontal discrimination of sounds (ears on horizontal plane)... not so good at vertical

2)Active Hearing
-animals generate the sound energy and detects the echos
-eg: bats (hear 9-212Hz, but use low freq in hunting because high freq fades fast), cetaceans (hear u100Hz, very accurate), birds etc

-wavelengths 10-20Hz
-rarely used, but homing pigeons can detect as low as 0.05Hz
-may be used to detect wind movement over mountains, wind of thunderstorms, earthquakes etc.
-the ability to detect electric currents
-found in all main groups of aquatic vertebrates (eg: platypus [mammal] has electric sensitivity on bill) and amphibian larvas

-2 types
i) Passive: ability to pick up currents by muscles of other individuals (ex: sharks)
ii) Active: production of electric currents whcih interact with environment. Used for hunting and orientation (much like echolocation). Body must stay fairly straight and rigid to keep appropriate current alignment and to minimize own muscle contractions (interference)
-some insects, birds, mammals, and fish have magnetic compounds in body cells (Fe3O4)
-good for orientation. Ability to obtain/maintain a direction relative to an external cue
-good for navigation. Focus on a goal, specific location toward which animal is moving. Animal capable of adjusting for changes/displacements
-may be used in migration
How do birds MIGRATE?
-may involve magnetoreception to guide them
-involves movement from 1 location to another with subsequent retun to th first. usually 2 different habitats
-eg: Von Frisch bee navigation and orientation
Discuss Von Frisch's experiment with bee navigation and orientation
-experimenters provided landmarks at various distances from hives (colored tents). Once bees recognized tents, they switched tents around. Found that bees had trouble finding hive when color sequence of tents was altered
-once bees were trained at a food source, landmarks could be removed/substituted, bees still could find food. Not completely reliant on landmarks, but had built in directional ability

-bees use sun as a compass, direction or polarized light on cloudy days. Have internal clock that adjusts for sun angle during time away from hive

-migrations can be small or large scale (definition dependent)
most birds seem not to navigae very well, exept pigrions and some sea birds (no land to follow, must use navigational sense
-genetic and fixed vs learned
-both present in birds
-eg: shearwaters (sea birds) adults leave breeding islands before chicks can fly. Juveniles return to same island 2-4 years later to breed (and never seen at island before this tim). Don't learn this migration from parents, something imprinted early in life or fixed
-eg: peregrines cross fasterned nestlings learned migration routes of their parents (partially learned)
What is the ULIMATE QUESTION of Migration?
-less then 50% of waterfowl return to breeding gournds after migration maybe because: hunting, predation, weather and disorientation, energy and saturation
1)If migrants want to minimize travelling time (ie: maximize migration speed how should they optimize their fuel loading and consumption of food?

2)"Continuous stopover scenario". Individual migrant can stop along the way wherever it wants to in order to eat and boost reserves

3)How to trade of accumulating reserves prior to departure vs. the need to stop and refuel

**being heavy reduces flight efficiency and increases predation rist**

**being too light risks starvation if conditions are unpredictable along the route**
Based on the OPTIMIZATION MODELS of MIGRATION, what are the 2 possible model scenarios?
Global variation
-each stopover patch has the same quality, and so food intake rate is predictable for the migrant based on the information it has in its current patch

Local Variation
-the quality of patches varies locally (it may be better or worse than the conditions experienced by the bird at the current stopover site)

**Feb.24 HO to see graph**
How do birds achieve OPTIMAL FLIGHT SPEEDS?
-in order to maximize flight efficiency, aerial migrants may need to adjust their airspeed as winds change. Eg: airspeed should be reduced in a tail wind and increased in a headwind to maintain same speed of travel over landscape

Optimal airspeeds may differ according to traits of each individual
-males, or dominant individuals may need to reach the breeding ground very early to get the best territories
-females may favour a slower speed to conserve energy for reproduction

**Feb 24/06 HO**
How do biological rhythyms become matched to the environment?
Synchronized behaviour and physiology
1)Direct response to external (exogenous) geophysical stimuli
2)An internal (endogenous) rythm that programs an animals behaviour
3)A combo of 1 & 2
Give 5 pieces of evidence for the ENDOGENOUS NATURE of the RHYTHYM
1)Rhythm is not synchronous with any known enviro factor

2)Animals are kept in adjacent cages under identical conditions display rhythms with different periods

3)Rhythm persists when the animal is moved from 1 part of the world to another

4)Rhythm persists when the animal is brought into a lab and held under constant conditions

5)Rhythm of one individual can be transferred to another by means of tissue transport
-an internal rhythm that programs an animals behaviour
-ndogenous rhythms tend to deviate over time (from exogenous cycles), if left isolated from normal environmental stimulus
night/dark cycle

Animals only have 1 biological clock
-can have multiple clocks with different periods within the same individual
-some clocks may underlie other clocks

eg: migration triggering
>brains pineal gland is sensitive to light only in fixed time window
>as daylength increases n spring, those cells get stimulated in their sensitive phase
>hormones released -> gonads develope and increase in size
>reproductive chnages and "migration restlessness"
What are the 4 types of Rythms?
-rev of earth


-29.5 day
-phase of moon

-365 days
-season of year
How does HIBERNATION occur?
-a detection of temp change by the hypothalamus triggers the body to go dormant

eg: crabs
-raised in eggs in lab
-young crabs kept in constant light/dark regimes
-no endogenous rhythm appeared...
-then a single cold temp shock caused the normal tidal cycle to appear
-temp is the zeitgever for the tidal rhythm
-are distinct from the physiological and behavioural rhythms
-clock is an oscillator, a self sustained time keeper (eg: a pacemaker is an oscillator that regulates some physiological or behavioural funciton)
-the function of the clock is not dependant on the expression of its behaviour
-clocks can be inherited, and rhythms can be influenced by imrinting and be passed on from mother to fetus
-clocks are relatively independant of temp
-clocks are relatively unaffected by chemicals
-clocks have limits to entrainment
Where is the anatomical location of the master clock in mammals and birds?
-hypothalmic suprachiasmatic nuclei (SCN) of the brain

Birds (& lower vertebrates)
-multiple oscillators interact to produce a stable rhythm
-there are 3 autonomons and anatomically distinct clocks located in the retina, pinal gland, and hypothalmic osciallator
What did Aschoff (1989) discover about Human Bio clocks?
-explanation for sleep requirements?
1)Direct response to short nights (responding to external, environmental stimuli)
2)Endogenous: rhythm alwys present even at Southern lattitudes, but Tinbergen just notied the rhythm when 'reed' from social pressures. Expt. males and female were isolated from any time source, females slept 1 hr more than guys, although trends were similar

-many human behaviours had a circadian or circannual hyhm
-grip strencth was stronger during the day
-time seemed to pass faster during the day then at night (by estimation of 10sec or 1 min intervals)
What is necessary for communication to occur?
-other individuals
-don't have to be the same spp
-difficult to interpret the receiver's behaviour (are they geting the signal?)
-in animal behaviour, there is no other operational criteria for communication othr than a change of behavoiur of the receiver (sender->signal->reciever)
What is EO Wilsons defintion of Communication?
-"an action on the part of 1 organism that alters the probable patter of behaviour of another organism in a fashion adaptive to the participants"
-emphasizes that nat sel has acted on the signal to increase its effectiveness
What is H Whiby's defintion of Communication?
"any alteratino in a reciever produced by a signaller by means of a signal"

-channel; the medium through which the signal is transmitted (ie: visual, vocal)
-signal; the behaviour (ie: posture, display, chem) transmitted by the sender
-context; the setting in which the signal is transmitted and recieved
-message; what the signal encodes about the sender
-meaning; what the reciever construes from the signal

*signal does not = message does not = meaning
What is Dawkins and Krebs 'RADICAL' interpretation of Communication?
'Manipulators' -> Signal (honest or deceitful) -> Mind Readers

-signals evolve with the motivation of being manipulative
-receivers are skeptical, try to pick up on honest cues, but may not respond to the original signal *arms race

-signals evolve differently depedning on the motivation and benefits of sender and reciever
-discreet, informative, efficient
2)Arms race
-flamboyance, energetically costly, not necessarily informative about true intentions

*diff between cues and signals
Whats the diff b/w a cue and a signal?
-unintentional signal

-intentional signal
What 7 Factors influence the design of a signal?
-max dist
-speed to travel
-how long it lasts
-how easily it is located
-detectability relative to background noise
-the info it contains
-whether it can be easily faked
1)Ecology and Environmental Traits

2)Physical Structure (Patterning) of he signal

3)Social Functions & motivation of sender and reciever

***These will be in detail in other FC***
Expt: Alberts measured molecular weights of chemical signals in diff social context
-heavy molecules vs. light moleculs: light evaporates quicker but travels farther and faster

Molecular Weight Effects:
1)Signal rise time
-speed of transmission (ie: sex attraction
-ie: time to reach maximum spatial range
2)Fade out time
-heavier compounds have slower fade out times (ie: scent marking)
*was discovered that weight of compounds increases with temperature of environment

Eg: Bird Song, auditory signal
-the denser the environment (ie: lots of vegetation) the greater the wavelength of the signal to make it through
1)Attenuation of sound waves
-high freq travel less far then lower freqs (b/c longer wavelengths)
2)Sound degradation
-ie: echoeing, mixing with ambient noises
There are 2 types of Patterning
-binary or digital
-on or off
-continuously variable (analog)
-eg: color variation in fish or crest of a bird

This allows for complex signals. Needed b/c:
a)Backup hypothesis
-components may enhance eachother and reduce ambiuity
b)Multiple Message
-provides different types of info about the signaller
c)Redundant and Non-Informative
-'arms'race' between sender and reciever leads to wasted or ignored signals
-Formalized Signals: ritualization, an evolutionary process by which signals become stereotyped
What is Darwins Principle of Anithesis?
-when an animal reverses it's intention, it reverses its signal
What is DISPLAY?
-a behaviour, that through evolutionary processes, has been modified and ritualized to serve a communication function
-typical intensity of the display (non-variable between individuals
-remember, not all reeated behaviours are displayed
-can be very dramatic and obvious
-can involve color (a 'display', behavioural control)
-else-> a 'signal'

-song: eg, birds, primates (arboreal) singing vole, fish (air bladder vibrations), chest beating of gorillas, chirpping o grasshoppers, beaver slap, etc.
-can be vocal or non vocal

-common in mammals, not so much in birds
-allogrooming/allopreening (in birds) used in courtship
-mouth biting/licking -> canids
-insects: bees follow movements of others with antennae

-usually not a display unless some behaviour surrounding exuding or depositing the chemical is ritualized
-eg: alarm chem in fish. Cues but probably not displays, maybe not even signals in the sense that they are meant for communications
Give 6 examples of the kinds of behaviour patterns and other responses from which displays are thought to have evolved.
1)Intention Movement
-action that is begun but aborted before it is fully complete
-eg: biting mouth opens and lips drawn during the normal process of biting (early steps are ritualized)

2)Ambivalent/Conflict Behaviour
-caught in between 2 behaviours, stuck in 2 conflicking postures
-eg: attack vs escape

3)Protective Response
-do whatever to protect self
-eg: monkeys scrunch cheeks to protect their eyes

4)Autonomic Response
-involves homeostasis activities
-eg: sweating

5)Displacement Activities
-behaviours performed out of context
-sign of frustration or stress
-behaviours directed to atypical objects
-eg: preening in duck courship

6)Redirected Attack
-beating shit out of inanimate objects
Why has evolution favored ritualization?
1)Ethological classic explanation
-ritualization reduced ambiguity and increased communication function (repetitive, stereotyped, exaggerated, moevments emphasized by coloration)
-eg: courtship and mate attraction... want the correct spps!!!!
-*trade of betwen ritualization and amount of info conveyed
2)Behavioural Ecology
-doesn't always pay the sender to by completely honest
-signals must be honest in order to be believed (related to cost and benefit of sending a signal)

Eg: if 'A' manipulates 'B' in a way beneficial to 'A' but detrimental to 'B', then 'B' should develope "sales resistence"
-start of evolutionary process could be same under both scenarios. 'B' looks for predictive cues in 'A' behaviour
-therefore can get a build up of ritualized signals over time, most of which 'B' no longer believes
What are 6 hypothesis for repetitive or multi channel signals?
1)To reduce ambiguity of the message

2)Escalation between sales resistence and advertising

3)to aboid stimulu specific fatigue (neural basis)

4)Stimulus summation: build up of stimulus effects

5)A way to assess an individual over time=> why signals are repeated

6)Redundancy maintains physiological arousal or hormone release etc.
What is the Message/Info actually in a display?
1)Observe chain of events
-eg: kittiwakes facing away display = reduction in aggressoin
-eg: grackles male approach female during breeding. Female withdraws unless male is performing head down display. Female then repells male or allows copulation, but never withdraws

2)When chain of events is not so clear, then do a probability analysis

3)Experiment with components of displays
-eg: use models - taxidermic (Tinbergen using stickleback decoys with redbellies
-eg: playbacks. Splice different song bits together, change frequency and timing
-eg: Singing birds, seeking or interactional. It ID's the individual as spp 'x' as a male in breeding condition
Why do vertebrates have a decreased number of displays?
-maybe no more needed
-maybe neurological constraints on processing more displays
-displays maybe not so informative anyways
-maybe content of info is increased
How can info content be increased?
1)Graded Signals
-maybe just a shift in intensity but can convey very different messages
-eg: ants, alarm chemical: at low conc, move towards signal; at med conc, workers to into alarm frenzy; high conc lasting 1-2min, digging behaviour

2)Vary the context of signal
>eg: Lions roar
-1) location isgnal when pride members are separated
-2) reinforces bonds within pride when together
-3) keeps other prides away (territorial signal)
-4) part of aggressive display in close encounters

-a display is used to change the context of subsequent displays or acts
-eg: if dos bow to eachother before they play, anything goes b/c not serious
In summary, what information is to be expected from displays?
1)Info about the size, age, strength of signaller in ways not open to cheating

2)Not expected to give away intentions except in co-operative interactions

3)May contain info about environment- food sources, presence of a predator, etc.
What are the Roles of Displays in Aggressive Contexts?
1)War of Attrition
-waiting game
-evenly matched contestants
-whomever displayed the longest or persisted the longest won

2)Game Theory
-ESS have a random persistence time (therefore non predictable)
-eg: Siamese fighting fish; early displays are very poor predictors of subesquent actions in the fight sequence

3)Asymmetric context
-2 individuals unequal in some aspect
-eg: fighting ability (body size, condition, or experience)
-eg: payoff asymmetry (one has more to gain from the resource. eg: hunger levels, desperation, etc.

4)Resource Holding Potential (RHP)
-pays both ind's to assess RHP before fighting... avoids physically costly fights
-signals of RHP should not be able to be faked by weaker individuals (> costs = > chance of winning... tough bluff)
-Displays of RHP involve: 1)Energy costs; and 2)Laws of Physics (smller an't match larger in some aspects [ie: pitch of frog croaks])

5)Aggressive contests
-graded stages of escalation
-fights should escalate more if he 2 competitors are more equally matched
-bluffing = displays to exaggerate body size. Limits on bluffing because at some point, the bluff will be tested with a real fight
What are badges of status?
-a visible cue of how dangerous a mother fucker is
-ie: a big black patch on chest

**it's not enough to look the part, or to be the part. It was determined that you need to be both in order to rise in status**

-some badges are correlated with age (eg: bald eagles, white head at 5yrs shows breeding status and dominance)
Do animals cheat?
-in low freq
-eg: Beau Geste effect
-illigitamate sender, like the boy who creis wolf, gives alarm calls so competition fucks off and gets prey
-false projection of another sp ID'ing signals and preys on duped males (ie: firefly's)
-illegitimate reciever - often predators (eg: bats can pick pu on some male frog croaks, therore frogs either stop or change pitch
What is the Beau Gaste effect?
-make seem like there are more individuals defending a territory then there actually is. used by some bird songs
-song repetoire may increase with male age and experinece
-may reliably signal fighting ability (therefore not Beau Geste) but not necessarily tru in al lbirds studied
-often predators
-costs: risk of injury or death
-benefits: value of resource (hunger level, RHP)

Solution to the game (ESS) Mar.13 HO
V=value ; C=cost
-1) if V>C, then hawk isESS, dove strategy can't invade a pop'n of pure hawks
-2)if C>V, then nither pure hawk or dove is ESS. Get a stable mixture, p=V/C
What is Maynard Smith's Bourgeois Theory?
**based on Mar.13 HO**

-play hawk if initial owner (eg: of territory
-play dove if you are intruder
-if it was modelled as 50% of the time owner vs 50% of intruder, then Bourgeois was ESS

a)Serious Fights
-commonly over mates because maybe only chance to pass on genes
-60% of male narwhals had broken tusks or fragments imbedded in wounds
b)Bourgeois Strategy
-respect for ownership
-when V is low or C is high
-the study of the biological basis (adaptiveness) and patterns of social behavoiurs in all organisms including humans
What is a SOCIETY?
-a group of individuals of the SAME spp organized in a cooperative manner
-reciprical communication transcending mere sexual activity is required

*note: some animals congregate but not as a society (ie: moths around light)
What are the 11 Degress/Traits of Society?
1)Group Size
-minimum is 2, but more than just mating involved
-should consider group size relative to pop'n size
-perhpas % of individuals found in a group is a better measure

2)Demographic Distribution
-age structure, reproductive rates, etc.
-eg: parental care or altruistic behaviour given by 1 individutal to another ="cooperation"
-behaviours directed according to differenct demographic subclasses
-functioning of society depends on proportions of the differrent group types

-wide range of communication can occur at close distances
-effectiveness of cooperative behaviours may improve with cohesiveness
-close spacing between individuals may indicate high sociability

4)Pattern and Amount of Connectiveness
-are signals directed preferentaially to certain subclasses or randomly to all within 'earshot'
>lower connectiveness, despite many individuals reached by the signal
># of ind's contacted per unit time by the average group member measure of connectiveness
>heirarchies and levels of communication
>signals pass to only subsets of the group
>members specializeing for different functions, therefore efficiency and coordination of the group can increase
-higher societies have high patterning, but not high connectiveness

-impermeable: few immigrants and little contact with other groups
-eg: baboon troop, only permeable when a new male takes over the group. Chimps have much more permeable groups
-consequences... gene flow, degree of relatedness etc.

-how much subgroups operate as discree units. Measure of complexity of the society
-eg:wildebeestes flee as disorganized mob when confronted by a competitor (compartmentlization low)

7)Differentiation Roles
-subgroups take on different roles within a society. Maybe more efficient to specialize rather than everyone being generalists
-eg: wild dogs. Role of hunder, Role of protector

8)Integration of behaviour
-measuring how well coordinated are behaviours among subgroups of a society

9)Info Flow
-total # of signals
-amount of info in signal
-rate of info transfer

10)Fraction of effort devoted to social behaviour
-% of time in day/% energy budget used for social interactions group activity
-eg: social insects approx 100% effort goes to colony except for self grooming and foraging for self, etc.

11)Minimum specification
-# of individuals that must be gathered together to see full behaviour repertoire of spp
What are the PREDATION factors considered when forming a SOCIAL GROUP?
-actually attracting predatory attention

-vigilance increases, *self herd effect
-*behaviour depends on location within the group
-eg: individuals on edge of gropu scanned more than those in the center
-dilution effect across more individuals
-confusion effect
-alarm calling
What are the FORAGING factors considered when forming a SOCIAL GROUP?
-fighting and competition because you must share energy supply
-take larger prey as a group
-defend prey better against other ompetitors

eg: competition between lions and hyenas
-lions less likely to successfully defend against hyenas compared to prides
-cooperative hunting techniques improve efficiency of prey capture
-eg: killer whales (15-20) hunt porpose schools. Group herding of prey and then each takes turn feeding while others stay back to prevent prey's escape
-eg: pelicans also group herd to feed
-eg: harris's hawk have flash and ambush technique
-eg: hyenas distract mother while another kills it's calf
Whats the difference between PRODUCERS and SCROUNGERS?
-social learning and information gathering about prey learning of dangerous food items
-get info about good food patches by monitouring where others go (puclic info)
-may lead to producer vs scrounger strategies in a group
-scroungers parastize producers efforts
-ESS occurs where payoff to producing = payoff to scrounging (Mar 15 notes)
What are 2 costs of Sociality?

Knowing this, why would Swallows hen be colonial?

-eg: cliff swallows
-mud nests packed into colonies
-rates of infection increase with colony size

1)Constraint of nest placement
-good nesting cliffs were rare

2)Using group information to improve foraging success
How big should groups be?
-proportional to how rich/poor resources are
-proportional to predation
-the minimum distance an animal routinely keeps between itself and others of the same spp
-maintained either by retreating or threatening an encroaching neighbor
-eg: a few fish in schools do not show this, bodies touch and shit
-eg: barn swallows 15cm, black headed gulls approx 30cm
-eg: sheep approx 1m and 60 degrees from nearest neighbor
-eg: humans body bubble, South Eur and asia = small space; NA = large space
-a min dist a pred is allowed to approach before an ind flees
-varies between spp
-a proximate cause of spacing but usually the ultimagte cause is centered on competition for resources

Individuals in a group may:
1)Defend large areas withall required resources
2)Defend point source with particular resources
3)Establish priority for access to resources
-an attribute of the pattern of repeated, agonistic interactions between 2 individuals
-characterized by consistent outcome in favour of the same dyad member and a default yielding response of its opponent rather than escaltion
-the status of the consistent winner is dominant and that of the loser subordinate
Does previous experience and interaction have an effect on fights?
-yes, it is easir to win when you are on a winning streak
Does Dominance = Aggression?
-middle ranked ind fights most often b/c ierarchy not clear cut
What are 2 forms of Dominance?
-one dom, bunch of subordinates, doesn't go further than that

2)Linear (ie: alpha, beta, gamma etc.)
-more common
-may be complex at the beginning when everyones trying to figure out who is top dog (ie: triangle relationship)
-unstable so tends to become linear over time

*once a group has ben set up, aggression decreases as signals increase to indicate dominance
If you are SUBORDINATE, why stay in the group?
-better than being solitary

3 Important Advantages
i)food increase
ii)social harassment decrease
iii)predation risk decrease

-make the best of a bad situation
-using alternate reproductive tactics (ie: sneaking, and you don't have to defend!! cha-ching!!!)
-put in time until they become dominant
Why do Dominants Tolerate Subordinates?
1)Not worth time and energy to chase them away

2)dominants also get direct benefits from a larger group
-eg: dominance in feeding piglets and milk gradient
What are the Economics of Territory Defence?
1)Spatial Distribution and Abundance of Resource

2)Population Density and # of Competitors
-an area that an animal learns and habitually patrols
-a defended area
-a space from which an ind, or group of mutually tolerant ind's actively excludes competitors for a specific resource
1)Floating vs Fixed Location
-eg: bitterling fish laying eggs on bivalve shells; display areas of fish are located above clam shells
-eg: male vertebrates (deer) defend moving harems of femals

2)Method of Defense
i) Absolute
-entire perimeter is constantly guarded and defended
-When Territores are relatively small (eg: fish, lekking grouse) or good vision or other distant sensing capability (eg: birds, raptors, etc.)

-resident only defends if is encounters another individual at close range
-eg: mammals... (bears)
-generational transfer from parent to offspring, therefor boundaries might be fairly stable for generations and not a lot of fighting # of encounters depends on pop'n density

3)Stability of Boundaries
-change with season, pop'n density, age or traits of the ind

-eg: Elastic disc model
>High Densities: breakdown of territory... dominance
>Low Densities: gaps between territories. Ind's no longer contact eachother

4)What is defended?
-Type A: large defended area containing resources for all the animals needs. eg: smalle insectivorous birds, small mammals, arboreal lizards, benthic fish

-Type B: large area for breeding but doesn't contain main food source. eg: nighthawks

-Type C: Small defended area around nest. eg: many colonial birds, flicker, insects (bees and wasps)

-Type D: pairing with or without mating (leks). eg: damselflies, ungulates, grouse

-Type E: defnece of roost or a shelter. eg: bat roosts, bird roosts (pigeons and starlings)
-food access increases
-renewable food sources can be harvested at a higher rate (nector flowers)
-food supply might be more stable
-predation risk may decrease
-disease and parasites decrease
-number of mates increase
-conditions for offspring increase
-eviction of previous owner
-maintenance costs (display = energy and time cost) (aggression = injury)
-time consuming to monitor and patrol area
-Type D territories don't have enough food
-decrease in the body condition (eg: male harem holders)
What are Ecological Factors that affect Cost/Benefit Tradeoffs?
1)Resource Quality & Distribution
-if food is poor quality (not enough calories) and sparsely distributed, then no defense (eg: Grazing Wildebeest)

2)Resource Distribution in Time
-ephemeral sources lead to a wide area needed to sustain self... therefore territory is not economical
-as predictability increases, maybe pays to exploit the same area over a longer time

-as # increases, costs increase

*see Mar.22/06**
What is the optimal Territory size?
-optimal sizze is variable depending on the situation but lies between a & b of a cost benefit curve
-at 'a', costs are minimalized
-at 'b', benefits are maximized
Consider 3 territories with low, med and high nectar levels. What would be the best defense strategy for each?
Low nectar levels:
-at low production, they abandoned territories... couldn't get enough to survive and pay defense costs

Med nectar levels:
-defense has huge benefits, so renewal rates make it worthwhile to build up a good supply

High nectar levels:
-defense not worthwhile because resource superabunant and not much depleted by competitors... plenty to go around
How is interspecific territory related with TERRITORY SIZE?
-as territory size increase, number of competing species increase
-knows habitat quality
-no territoriality, but there is competition

**see Mar.24 for the graph***
-monopolization by an agressive individual
-floaters are comletely excluded from territories
**See Mar.27 notes**
1)Ind differ in competitive ability and some exclude others

2)Fitness gain in 2 habitats is not equal

3)Some ind may be excluded altogether (floaters)

4)less variation in #'s in a good habitat than a poor one

5)Can lead to reulation of pop'n size if # territory is limited
-fills good habitat first then spills over to shitty ones... birds in shitty place are usually birds of shitty rank (ie: floaters)
What are the strategies of Floaters?
1)sneak resources from territory owners (eg: matings, food, etc)

2)Patrol wide area... search for vacancy

3)waiting to gain in age and experience

4)eventually... might get a territory

5)disperse if local prospects aren't good
-passing genes to next generation. 2 ways:
1) Survival: Darwin's natural selection

2) Reproduction: getting mates, increasing number of offspring
Are females picky about who they hump?
-they won't just hump first dude
-they go around and judge dudes, pick the best and then hump the hell out of them
(eg: peacock, highest eyespot in dudes = lots of pussy)
Give 3 examples of male-male competition in peacocks.
1) Train used in agressiveness. Raised in response to other males, not shivered as in bowing display

2) High status male have more elaborate trains

3) High status males gain more matings

*However, no corelations between indicators of dominance and train morphology
What are 6 reasons why male ornaments have evolved?
1)Pleiotropic gene effects

2)Selection of ecological sex differencese

3)Males being unprofitable prey for predators

4)Male contests

5)Female choice and mating preferences

6)Wallace: Dichromatic animals
-females have good sense (ie: choose most vigorous males [nat sel])
What are 6 reasons female why female preferences for male ornaments evolved?
1)Fisherian self reinforcing selection
-AKA 'Run-Away' Selection
-by chance you get a male that has a special trait (ie: long tail)
-in the pop'n are females who like that special trait
-so he gets all the humping, and next gen carries that trait

2)Indicator mechanisms
-indicates 'health' or viability of the male (something that is heritable)
-therefor choice benefits the females fitness
-eg: Hamilton ZEck hypothesis states that color may be a signal of parasite resistances

3)Selection for spps recognition
-hard to experiment

4)Direct phenotypic benfits to choosy females
-male birds often courtship fed the females
-the bitch chooses based on male care

5)Selection of the sensory system in other contexts (sensory bias)
-Darwin: goodtaste (male beauty)
-a novel trait but somehow attracts mates attention
-uses tricks to attract shit in different ranges
-organs may evolve just i ntime for reproduction

6)Advantages in the timing of reproduction (mating synchronization)
What are 3 forms of male-male competition?
-quick searching and location of mates

2)Endurance with Lek or Harem

-usually favors larger sizes of males and sexual size dimorphism, sometimes small so they can 'sneak'
-eg: roaring contest, prallel walks, physical contest
-eg: sperm collection. damselfly has bristles in his dink which scoops out competitions sperm while he shoots his in. Then leaves some protein in there so no more can get in
What is INFANTICIDE and why would a male commit it?
-if by killing a females offspring, a male can initiate a new breeding cycle sooner
Why is it usually males that compete?
1)Bateman Gradient
-more mates = more offspring

2)Triver's Parental Investment Theory
-PI=investment in an offspring that reduces a parents ability to in vest in other/future offspring -vs- parental care = any behaviour that improves the survival of an offspring
-when 1 parent invests more than the other, it should be choosy beause the cost of making a mistake is large
-eg: female mammals lacation goes on for weeks/months/years, males can desert and leave entire cost to females

3)Operation Sex Ratio
-more abundant sex will compete for the rarer sex
-eg: females input high energy, therefore have short lifespans, therefor skews sex ratio, therefore males compete for females
What are the 2 major types of Mating Systems?
-male and female have 1 partner per mating/breeding season

2)Polygamy: either male or female has >1 mate in a breeding season

2a)Polyandry: female with >1 male

2b)Polygyny: male with >1 female

2c)Promiscuity: both male and female mate with multiple partners with no pair bonds

2d)Polygymandry: male and female mate with multiple partners but form pair bonds in a group of co-breeders (eg: cooperatively breeding spps)
When should Polygamy be more common?
1)one sex is predisposed to assume most of the parental care(ie: if difficult to leave because of duties) (eg: lactation in mammals)

2)Parental care requirements are minimal
-ie: 2 parents not needed to rear young [precocial(leking spp of grouse) -vs- atricial spp(2 parents needed to feed young)]

3)Super abundant food resources enables a single parent to provide parental care
What are the economic defendability of resources (mates) dependant on?
Spatial Reason
-it is difficult to monopolize resourses that are uniform in space
-patchy (clumped) resources may be defended by territorial individuals

Temporal Reasons
-if all females are sexually receptive at the same time and for a short time, it is hard to monopolize

Spatial and Temporal clumping influences operational sex ratio = the ratio of gertilizble females to sexually active males at any given time

-if evenly distributed, then little polygamy potential
-if patchy distributed, then high polygamy potential
-best strategy for both partners because a single parent can rear few or no offspring alone
-biparental care is not needed to raise offspring
-partners stay together because of constraint
What is the purpose of doing a MATE REMOVAL EXPERIMENT?
-to see which sex is better at rearing offspring
-females are needed, dudes sucked
-but in woodpeckers, dudeds did great
Short term
-productivity of the first brood that is left with the deserted partner

Long Term
-provisional effort and survival of single parents
-chance of repairing with same partner

*Trends (in woodpeckers)
-0 males abandoned, low # of fems did
-males reared 85% fleding success, fems reared 41%
-nestlings died early with fem than with male
-males defend resources that fems need
-males compete among themselves over subdividing and defending resources
-fems choose mates based on male quality, but largely on value of resources he defends
-incoming fem must decide whether to mate with; 1)An already paired male [share resources and care], or with a; 2)unpaired male on a poorer territory [gets all, but may suck]
When will males get super choosy?
-if males invest lots in monogamous situation, then they may become choosy or adjust investment based on female quality or assurance of paternity

Eg: Dunnock breedin system
-females do least well under polygyny because they must share male care with other females
-females do best with polyandry (many males help feed the chicks)

*mating system in Dunnocks partly depends on ecological factors
-fems defend small territory when food is supplemented
-size of males territories do not change
-a given male can overlap more female territories therefor potential for polygyny increases, polyandry decreases

What 6 factors are characteristic of PARENTAL CARE?
-any parental behaviour that increases the reproductive potential of an indiv's offspring

1)Making nests, burrows, territories, etc.
-parly mating effort in addition to care (try to impress)

2)Production and provisioning of gametes
-size/quality of eggs (eg: amount of yolk)
-male nuptual gifts

3)Care of eggs
-brooding/guarding (fish, insects, some amphibians, crocs etc)
-incubation (birds)

4)Guarding offspring
-after hatch

5)Provisioning of offspring
-providing other food

6)Care after nutritional independence
-long lived social spp
-teach offspring, help acquire territory
Should the male or female desert the offspring?
-answer may depend on what the mate does in response. Stay or leave?

3 Possible outcomes:
1)both desert
2)1 stays, other deserts
3)both stay

Solutions depend on:
-the survival of the offspring under each scenario (even if some offspring die if 1 prent leaves, that parent may increase repro success if it can find another mate)
-which sex deserts may depend on which has the first opportunity (it is typically the male, since he doesn't lactate, or gets sore from giving birth... hahaha!)
How do parents direct their care?
Trivers: Parent offspring conflict
-parents are related by 0.5 to each offspring
-an offspring is related 1.0 to itself and 0.5 to siblings
-therefore offspring may demand 2x the amount of care that the paretn is willing to give
Why would parental care conflicts arise as the offspring age?
1) Actual costs to parents increase
-food and energy increase as offspring increase in size

2) Benefit to offspring plateaus
-becomes more self sufficient
-after time, parents would do better investing in a new/younger offspring
-leads to weaning conflict
-signal need
-open to cheating though (damn con artists)
-energetic and predation costs might keep begging honest
-some signals may be honest (ie: gape color, weight, etc)
-because parents are r=0.5, they are selected to invest equally in offspring (when all else equal)
-each offspring will demand more to self (r=1) than to siblings (r=0.5)
-parental favouritism may occur if some offspring are better quality than others
-parents may observe outcomes of sibling aggression to see which is worth investing in
-parents start with optomistically large brood
-if food is unpredictable, a few disadvantaged young can be quickly killed off and therefore brood size gets adjusted according to food availability
-or parents can let this happen naturally
When should animals fight?
1)Pop'n will tend to be all fighters if reward is far greater than cost (everyone wants a piece)
-fighters decrease as costs increase

2)Pop'n will be a stable mixture of fighters and pacifists if rewards are less than costs

3)A popn pf 100% pacifists is not stable because a rare fighter genotype will at first meet only pacifists in the population and take advantage of them
-being a fighter only becomes disadvantagrous once it starts encountering other fighters
-reduction in predator pressure by improved detection or repulsion of enemies
-improved foraging efficiency for large game or clumped ephemeral food resources
-improved defense of limited resources (space, food) against other groups of conspecific intruders
-improved care of offspring through communal feeding and protection
-increased competition within the group for food, mates, nest sites, nest materials, or other limited resources
-increased risk of infection by contagious diseases and parasites
-increased risk of exploitation of parental care by conspecifics
-increased risk that conspecifics will kill one's progeny
1)more copulations, more sex partners, or both
2)more copulations with estrous females
3)more sired offspring

4)more harassing of ohters while others were copulating
5)fewer offspring died following birth
6)longer life span or reproductive years
1)more offspring born
2)more offspring survive
3)earlier age, or earlier in season, at first conception

4)short birth intervals
5)reproductive suppression of other females
6)more copulation and/or more bonding to males
7)more harassing of couples while copulating
8)longer life span, longer reproductive career
What are some mechanisms in sexual selection? 7
1) Scrambles
2) Endurance rivalry
3) Contests
4) Mate choice
5) Coersion
6) Sperm competition
7) Infanticide
What do Females choose males by?
1)Good Taste (Darwin Fisher)

2)Good sense (Wallace)
-good genes
-good resources (genetic and non genetic)

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