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Dr. Underwoods animal behavior final Kutztown University fall 2025
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What is migration?
non-random directional movement that is predictable over time
-animal continues movement until suitable resources are encountered
Why and how do Wood Thrush birds migrate?
-breeds in Eastern PA/ US during summer
-leaves to escape winter
-spends 8 months in central america
-goes back north to escape competition for food and nest sites, exploit seasonal abundance of food in north, longer days for foraging= better reproductive success
What are the costs of bird migration?
-lots of energy, especially with storms; greater risk of death
-predation
-collisions with windows, lighthouses, towers…
What are neotropical migratory birds?
breed in northern temperate areas and winter in central/ south america
What are the benefits of the north in the summer and the south in the winter in terms of migration?
North- breed in summer
more food
less nest predation
less territory competition
better reproductive success
South- winter
reliable food
stable climate
higher winter survival
What are the benefits of being a permanent northern bird? (Black-capped chickadee)
access to resources
no migration
earlier nesting
higher reproductive success
What are the costs of being a permanent northern temperate bird? (Black-capped chickadee)
lower winter survival
What are the benefits of being a permanent tropical bird? (Golden-Hooded Tanger)
higher survival
access to resources
no migration
What are the costs of being a permanent tropical bird? (Golden-Hooded Tanger)
high nest predation
high competition for territories
How and why do Green Sea Turtles migrate?
Feed in waters off Brazil
More food + energy
migrate 1800km to Ascention islands to East
isolated beaches
few predators
reproduce and nest here
How and why do Gray Whales migrate?
one of the longest mammal migrations
feed in waters off of Alaska
more food + energy
Migrate 10,000km to Mexico
calve in warm coastal waters
less predators
How and why do Monarch Butterflies migrate?
Breed in US and Canada
more food for caterpillars; milkweed
Migrate 4,000km to Mexico
mtns to 10,000ft
overwinter in fir trees
shelter, prevent freezing, but not too hot
reduce metabolism- low energy use
Come back to TX/ Louisiana in spring to lay eggs
next gen comes back up north
How is climate change affecting Monarch Butterflies?
some now overwintering in southern US
going to higher elevations
80% reduction in our population
99% reduction in Cali/ western groups
Who are usually the choosers in mating?
females
who are the competitors in mating?
males
What is fighting/ aggression in securing copulations?
males fight steryotyped contests
-ex: fencing with antlers/ horns, head butting
Mates with larger body size and/ or weapon wins
Con: injury
what is dominance hierarchy in securing copulations?
dominant males inhibit subordinate males from mating
-aggressive threats
-disrupting mating
dominant has most copulations
-subordinate males mate our of sight of dominant males (surreptitious)
Are low ranking males condemned to celibacy?
no, just need to use alternate mating tactics
-male mating strategy is contingent upon competitive ability
High- large body size → compete for dominance
Low- small body → use “sneaker” strategy
What are alternative mating strategies/ benefits for large males?
-male body size and call intensity related to mating success
-large males with loud calls succeed in attracting females
What are alternative mating strategies for small males? (sneaker strategy)
adopt satellite behavior- intercepting females as they approach calling males (sneaker strategy)
intrasexual selection
what is mate guarding in terms of ensuring sperm success and what animal does it?
Amphipod- phylum Arthropoda, subphylum crustacea
Crustaceans mate for a short time after molting and do not store sperm
-male grabs female before mating
-rejects female if already mated
-if accepted, holds until she molts and sperm is transferred
-after mating, male carries female to prevent other copulations. Guards female longer if other males are present
What is sperm competition in terms of ensuring sperm success?
competition to fertilize eggs from more than 1 male mating with a female
-influenced by health of sperm, # of sperm, order of mating
what is sperm precedence?
timing of copulation relative to other males will determine success in fertilizing eggs
First male advantage
Last male advantage
sperm comp. favors males with adaptations to increase fertilization success
First male advantage
1st male to mate fertilizes most eggs
Last male advantage
last male to mate fertilizes more eggs
what is higher ejaculate volume in sperm competition?
testes size correlates with # sperm
-higher sperm= higher probability of fertilization
Gorillas: female rarely mates with >1 male
-smaller testes
Chimps: Female regularly mates with >1 male
-larger testes
what is frequent copulation in sperm competition?
dominant male guard (cant read word in notes? &) copulates frequently (Bighorn sheep)
-if subordinate chimp copulates, dominant male engages in “retaliatory copulations”
-copulate with that female even more than before
-may displace sperm by volume and sperm from last mating more likely to be successful (last mate advantage)
what is sperm displacement in sperm competition?
(penis of damselflies has backward hairs)
-scoop out sperm of previous males
engage in behaviors to stimulate female to eject previous male sperm
ex: dunnocks
what are repellents in preventing further mating?
male transfers anti-aphrodisiac pheromone with sperm
-female repulsive to future males
parasitoid wasps use scent to find offspring
what is a copulatory plug in preventing further mating?
thick secretion that blocks female reproductive tract
-rodents, snakes/ lizards, marsupials, bats, some primates
Possible functions:
-prevent mating, ensure sperm retained
-sperm transport; gradually released
-scent mark
common garden spider- very effective
what is sexual interference in preventing further mating?
< fitness of rival male by reducing mating success
salamanders produce sperm packet that is deposited on bottom of pond (spermatophore)
-female picks up and transfers to cloaca
spermatophore transfer interference
-rival male gets between courting of male/ female
-rival male deposits spermatophore to get female to pick it up
what is pseudo female behavior in sexual interference?
act like female to get rival to release spermatophore
what is infanticide in sexual interference?
kills competitors offspring
-outside male usurps resident male of a troop
kill offspring
bring female back to estrous (reproductive condition) so new male can mate
only kill unrelated offspring
(langur monkeys)
what are the direct benefits of female choice of a mate in intersexual selection?
material benefits
1.) sufficient sperm
2.) resources- food, nest
3.) parental care
what are the indirect benefits of female choice of a mate in intersexual selection?
benefit offspring
1.) good genes
2.) compatible genes
what is an example of female choice for sufficient sperm? (sato and goshima 2007)
store crabs
compared female preference for unmated vs previously mated (4x) males
-depleted sperm b/c previous mating
female prefers unmated- higher sperm
-chemical cues
important b/c female can choose from distance to avoid being caught/ held by male
what are nuptial gifts as resources in female choice of mate?
male provides nutrition/ resource
gift can:
> male mating success (balloon flies)
> number or fitness of offspring
ex: katydid- large spermatophore gives nutrients for female for eggs
arctiid moth male gives alkaloids from plants in spermatophore
australian red back spider lets female eat him, copulate 2x as long, fertilize more eggs →female less likely to mate
what are territories as resources in female mate selection?
(house wren)
secondary cavity rester- doesn’t build cavity for nest
males often defend >1 nest site
-nest limited supply
added nest boxes to territories (Eckerle and thompsoon 2006)
female preferred > nests
predation high, female can move to new site
what is parental care as a resource in female mate choice?
females prefer brighter red males (house finch)
put into captivity, controlled carotenoids in diet (red pigment)
must forage on food with high carotenoids while mating
bright red foragers indicates parental ability and red males fed chicks most food
what are good genes as an indirect benefit of female mating selection?
assess indicators of good genes
does tail symmetry affect mate choice? (moeller 1990-1992)
-female prefers long symmetrical tails
male barn swallow tail influenced by mite load
good genes for parasite resistance- healthier, better preening ability
whats an example of good genes in female mate choice for the American pronghorn?
male has harem of females (group of females the male defends for exclusive mating access)
-male provides no resources or care
why do females join?
-females prefer males who maintain “zone of tranquility”= area free from intruding rival males
-indicates male physical health/ vigor
mating success for males depends on # females in harem
what is compatible genes in reference to female mate choice?
offspring fitness may depend on fitness bt/wn male/ female genes
Major histocompatibility complex genes (MHC)
-important for immune response
-male house mice prefer diff MHC genes
-via odor cues/ chemical access to compatibility
why do animals choose compatible genes in terms of avoiding inbreeding?
1.) too much homozygosity
higher risk of genetic disorders (recessive) or lethal recessive alleles
2.) increase offspring resistance to disease with heterozygosity
what are indicator mechanisms?
traits indicate good genes
wha is runaway selection?
traits with no real benefits
-females prefer traits that can be passed over time
-male offspring more likely to have trait, pref/ trait linked
-increases selection for trait that may result in exaggeration of trait beyond natural benefits
-irish elk- massive antlers (extinct)
what is sensory bias?
traits that stimulate existing bias in female sensory system
-female prefers traits in males b/c linked to prior bias of females sensory system
Satin Bowerbird
color of females food used by males
-likes blue, males collect blue objects
tested foraging preference in females, they preferred blue food
what is the handicap principle?
-females prefer males that reduce survival- will have good genes
-male can survive despite handicap, female prefers male with largest handicap
ex: female peacocks prefer long tails/ eyespots
-foxes prey on males with lowest mating success
-male with long tail/ their offspring had higher success
-had better genes despite handicap
whats an example of runaway selection with sandflies?
-females choose mate over vertebrate host
-males with most matings called attractive, but didnt survive better; female didnt lay more eggs
-attractive male offspring had higher mating success but no actual better genes in particular
what is cryptic choice?
choice after copulation
cant see choice
female mates with >1 male, chooses which sperm fertilizes egg
males produce spermatophore
-tested males with diff quality by manipulating food access
-female chooses based on nutritional status
-manipulated female ability to control sperm (males mated with dead female)
more transfer in alive females for fed males. female controls sperm transfer
what is parental care?
parental investment
-expenditure on parents that increase offspring survival but decreases ability to invest in other offspring
-costly to parents and siblings; especially mothers
ex: desert spider- lets offspring feed on body
=matrophy
what is the cost of producing sons according to the Finland human pedigree data from the 17-1800s?
looked at offspring born after elder brothers
-survival is the same, decreased lifetime reproductive success
female cannot invest as much in next offspring
why do animals invest so much in sons?
males have higher reproductive potential
-can mate many times= many offspring
-females can only have certain number of offspring at a given time
what is the pattern of parental investment?
parents act to maximize lifetime reproductive success
-not each reproductive event
how do males and females decide patterns of reproductive investment?
1.) how much to invest in reproduction vs own growth/ survival
2.) how to allocate resources to offspring
-creates conflict between parents, parents/ offspring, and siblings
what is scrabble competition in sibling rivalry?
fighting to compete for parental resources
ex: domestic piglet fights for treats
-largest retains possession of food
-smallest fights for position- uses teeth as weapons (canines and 3rd incisors)
-if displaced, may not survive
what is siblicide?
killing sibling to obtain more parental resources
-more common in birds
ex: herons, pelicans, raptors
Why allow siblicide?
pelicans lay 2 eggs but almost always raise 1
insurance in case of:
-egg infertile/ doesnt develop
-food supply is unpredictable. If food is abundant more offspring than normal can be raised.
what are patterns of care in vertebrates?
Almost no care: amphibians, reptiles, fish
Mostly parental care: mammals, crocodilians, birds
Mammals- females provide most care, internal gestation + suckling of young
Birds- mostly biparental, require amount of food that one parent cannot provide
what are parental allocation of resources?
often males and females treated equally
Exceptions:
-brown songlarks- males are larger to attract more mates and have more reproductive success
-females feed offspring
-male nestlings grow faster and heavier, fed more/ higher quality food
males have higher reproductive potential, parents invest more
how does longevity/ reproductive strategy influence resource parental allocation?
birds tested by making feeding young more costly
-can trim parents wing or attach weight
-long-lived seabirds= dont incur costs- feed chicks less
-short-lived songbirds= feed chicks same- incur costs
short lived birds current brood of offspring makes up a larger percent of their lifetime reproductive success than long-lived birds
what is certainty of parentage in parental allocation of resources? with examples
males are less certain of parentage
bluegills- large fish will defend a nest/ care for offspring= parentals
small bluegills wont defend nest- sneaker strategy
Ex: parental nesting males exposed to sneaker males. Sneaker held in a container so it couldnt fertilize eggs
-parentals provided less care to eggs when sneaker present, but no difference in care of fry after hatching
-males can identify own offspring by chemical scent
social monogamy
birds- 90% monogamous
-10% genetically monogamous
most seek out extra pair copulations (EPC)
-no pair bond
What are the male benefits and costs of extra pair copulations?
benefit: higher reproductive success
costs: energy/ time
-cuckoldry- female has other male’s offspring
what are the female benefits and costs of extra pair copulations?
benefits: all eggs fertilized, higher quality genes, genetic diversity, resources
costs: reduced parental care
whats the example of extra pair copulations with purple martins?
old males produce avg 4.5 chicks with own mate and 3.6 chicks through EPC
What is polygamy?
pair bonds with >1 individual
whats the difference between polygyny and polyandry?
1.) polygyny- male has >1 female mate
2.) polyandry- female has >1 male mate (rarer)
describe polygyny
less or no parental care
more competition for resources
Why accept?
a.) polygyny threshold hypothesis
-female chooses a mated male IF quality of resources offsets cost
GREATER success as 2nd degree female on HIGH quality territory than monogamous female on LOW quality territory
whats an example of polygyny threshold hypothesis with the red winged black bird?
-if quality equal, female chooses unmated male
-if mated male territory is superior, female chose mated male
-females have more offspring in polygamous matings (graph on handout)
wha is the polygyny threshold?
point where there is no difference in reproductive success (PT on graph)
higher quality territory/ benefits must be greater than threshold for female to choose polygyny
what is the sexy son hypothesis?
-good genes for offspring offset polygyny
-produce sexy sons to have many grandchildren
pied flycatcher
-followed reproduction success of sons of females that chose monogamy and polygyny
found NO DIFFERENCE in reproductive success of sexy sons, no support for hypothesis
what is female defense in polygyny?
male defends female
when females are gregarious (live in groups)
-cooperate to hunt/ detect predators
-specific birthing grounds
ex: elephant seals
what is resource defense in polygyny?
males defend resources needed to reproduce: nests, food
-common in marshes/ grasslands
-bc resources are unevenly distributed/ clumped/ patchy
ex: marsh when
-male mating success related to vegetation cover
-54% cover= unmated (worse habitat)
-80%= monogamous
95%= polygamous (best habitat)
what is Lec polygyny?
males defend symbolic territories with no resources
occur in Leks (happens in birds/ bugs- midges)
-Lek= mating areas w/ those territories
-common in grouse and pheasants
-10% of dominant males get copulations, usually largest, defend central most territory
why did Lec polygyny evolve? in the male perspective
attract females, safer from predators,
hot spot hypothesis= locate Lek where > likely to encounter females
“hot shot hypothesis”= subordinate males will raise mating success by being near highly successful males
(sneaker males)
why did Lec polygyny evolve? in the female perspective
groups of females
safer from predators
avoid harassment from males
large group of males to choose from/ facilitates mate choice
what are the male/ female perspectives of polyandry?
Male perspec:
male has lowest reproduction sucess
-cant ensure paternity
-limit access to females= limit reproductive success
Female perspective:
-maximise reproductive success
only works if she gets male care/ monopolozes male parental care
what is altruism?
reducing your fitness for benefit of others
what is kin selection?
reducing your fitness for benefit of offspring
are alarm calls costly?
yes but only for terrestrial predators, not arial
what is reciprocal altruism?
does not always involve kin
actor incurs some cost helping recipient who returns favor later
selected for if net fitness go into altruist when favor returned
not dependent on kin
diff types- grooming, sharing food, alarm calls in groups, mixed spp with birds
what are the dependent factors of reciprocal altruism?
1.) high probability for repayment
2.) altruist and recipient recognize each other
-social spp. in groups
-indiv spp that live long
3.) benefit to recip > actor
what is an example of reciprocal altruism in vampire bats?
live 18 years in small stable groups
-feed on mammal blood- cows/ horses often
-1 meal sustains 48 hrs
-unsuccessful 1/3 nights of foraging; need sharing
likely that bats recognize each other; high prob of repayment
how do vampire bats share food in reciprocal altruism?
-needy bat starts licking fur of nearby successful bat
-needy licks successful bats lips
-actor regurgitates food
¼ of blood meal
recip- 12 hours of E
more beneficial to recip than cost to actor
how do vampire bats choose who to share food with?
recognize them- mates or relatives
greater opportunity for reciprocation the longer they know each other
in graph- r= coefficient of relatedness 0-1
kin share but not necessary
what is brood parasatism?
bird lays eggs in other bird’s nest and leave all care of offspring to foster parent/ host
what is a facultative brood parasite?
sometimes lays eggs in other nests, usually conspecific, ducks
what is an obligate brood parasite?
always a parasite, never builds own nest
rare- 1%
can lay more eggs than typical nesting bird
studied the most
what are some examples of an obligate brood parasite?
Common Cuckoo bird (host specialist)
brown headed cowbird (host generalist)
what is a host specialist?
only lays eggs in nest of 1 species/ a couple in an area
what is a host generalist in obligate brood parasites?
lays eggs in several spp nests
brown headed cowbird
what are the costs of parasitism for victims of cuckoo birds?
-paratatizes little song birds
-eats/ removes egg from host
-when hatches- push other eggs out
-needs more food (4x)/ space bc of size
-sometimes hatch together and throws out host’s chick
-host never raises own offspring
-total loss but rare (<5% get parasitized)
what are the costs of parasitism for victims of cowbirds?
-don’t evict hosts chicks, can throw an egg
-evolved to hatch before hosts chicks
-not total loss/ lower cost but reduce offspring for host
-cowbirds abundant- high freq.
multiple parasitism: multiple cowbird eggs in nest, they compete with eachother still
how do birds protect themselves from cuckoos and cowbirds?
1.) nest defense
2.) egg burial and nest desertion
3.) egg ejection
what is nest defense in brood parasite hosts?
can recognize eggs, cover nest. can still be attacked later
what is egg burial and nest desertion in brood parasite hosts?
build new nest on top of old nest BUT cowbird can keep coming back and laying eggs on top of new nests
-can leave nest BUT cowbird can find new nest
what is egg ejection in brood parasite hosts?
-pull out foreign egg, needs to be able to recognize
-if cowbird or cuckoo comes back, will do it again
what is cuckoo egg mimicry?
cuckoo will match look of hosts eggs
evolutionary arms race
-series of adaptations and counter adaptations between parasite/ host spp
-can also be with predator/ prey
what is a mating system?
social bond between males and females for reproduction
male and females at odds
what are the male and female restraints for reproductive success?
Male:
-access to females
-greater success by mating with more females
Female:
-parental care
-higher quality sperm/ good genes
-resources: food, nest sites
each try to maximize own success