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Sexual Selection
The difference in mating or fertilization success that arises from contests among males for mates and female preference for specific males
Sexual Selection Reality
Preferences in one sex drives the evolution of appearance & behavior in the opposite sex
What does sexual selection explain?
the existence of characteristics that do not confer a survival benefit, heritable
3 Hypotheses for Sexual Selection
- good genes
- direct benefits
- arbitrary choice & runaway selection
Good Genes Hypothesis
Elaborate plumages/displays signals genetic/physiological quality
Handicap Hypothesis
a male that can support a costly and unwieldy ornament is likely to be a vigorous individual whose overall genetic quality is high
Direct Benefits Hypothesis
female prefers male that provides tangible resources that increase fecundity; not necessarily genetic benefits
Arbitrary Choice & Runaway Selection Hypothesis
Sexual selection evolves through arbitrary choice and runaway selection for aesthetically pleasing features with no evolutionary advantage (can be heritable)
Results of Sexual Selection
sexual dimorphism
Why is sexual selection usually shown in males?
the sex that has the lower reproductive burden is the one in which sexual selection primarily acts
Pharalopes
females show sexual selection driven traits and lay many clutches, leaving males to incubate and care for young
Types of Displaying
- Courtship Building
- Vocalizations
- Lekking
Courtship Building
ex: male baya weavers build nests that females judge, selecting the male with the best quality nest
vocalizations
calls and songs themselves are often displays that have been selected for over time via sexual selection
lekking
The gathering of males in an area for the purpose of displaying to attract females. (no resources aside from being a place to display)
3 Hypotheses Behind Lekking
- Hot Spot Model
- Hotshot Model
- Female Preference Model
Hot Spot Model
males gather at sites where they are most likely to encounter roaming females; good positioning offsets the cost of competition
Hotshot Model
males gather around experienced, attractive, or dominant individuals to increase their chances of being noticed
female preference model
- females prefer to visit large clusters of males over small clusters or solitary males
- allows females to make safer and more efficient comparisons between individuals
kin selection in lekking
birds can benefit via increasing the fecundity of genetic relatives, even at the expense of their own reproductive success
Ruff Lekking
3 types of makes:
- independents (display at leks)
- satellites (track wandering females and are recruited as display partners)
- faeders (look like females and solicit copulations)
sex determination in birds
all birds have 2 sexes; determined at fertilization
female ability to control sex ratio
female birds can bias the sex ratio of their offspring based on their physical condition or needs of parents (unknown how)
Bilateral Gynandromorphy
Create offspring with half body of a female and half of a male
What are the 4 sexes of white-throated sparrows?
- white striped (male or female)
- tan-striped (male or female)
white-striped white-throated sparrow
aggressive, tuneful, poor parent, mates with tan-striped
tan-striped white-throated sparrow
less aggressive, better parent, mates with white-striped
Bird Sex Hormones
testosterone (aggression & courtship) and estrogen (parent care)
bird testes are internal
- attached to the dorsal body wall of the back of the kidney
- sperm evolved to survive high temps
do birds have penises?
ancestral trait, waterfowl and some other taxa have penises, but most do not
How do birds mate?
cloacal kiss
cloacal protuberance
sperm are stored at the terminal end of the vas deferens (seminal glomus), and this creates a swelling
how often are bird copulations?
rapid and frequent during egg laying; up to hundreds per clutch of eggs
bird sperm competition
- sperm from multiple males can internally compete to fertilize eggs
- females can eject sperm though generally most recent is most likely to fertlize
Male Adaptations for Sperm Competition
- larger testes, sperm stores, & frequent copulations
egg development
ovum -> yolk -> germ cell (20 hours)
how many ovaries do birds have?
1
Parthogenesis
birds cannot reproduce via unfertilized birth due to sex determination method
What is the purpose of eggs?
provide protection and containment for all of the nutrients necessary to build a hatchling from an avian embryo
What do eggs require?
external heat, periodic turning, and defense from predators and brood parasites
How is egg shape influenced?
stronger flight = more elliptical/asymmetrical eggs
Analogous to human placenta
chorioallantoic membrane (exchanges O2 and CO2)
Proximate effects on clutch size
- huge metabolic costs
- 20% energy transfer efficiency
Capital breeders effect on clutch size
stored resources are used to produce eggs
Income breeders effect on clutch size
- based on caloric intake
- need to acquire calcium
Ultimate factors that affect clutch size
- shaped by natural selection (varies from 1 to dozens)
- depends on ability to incubate eggs/care for fledglings
monogamy
predominant avian mating system (~90%) with 2 parents involved in the raising and care of offspring (typically when male help is essential)
serial monogamy
monogamous for a single breeding season or nesting attempt, typically practiced by songbirds (migration mortality)
lifelong monogamy
common in long-lived birds and occasionally short-lived species and resident birds
divorce
Some birds will leave their pair bond for higher-ranked/better-adapted individuals (infidelity is incredibly common)
Extra Pair copulations (EPC)
A mating by a male or female with someone other than their primary partner in a socially monogamous species, 76% of monogamous species (common in Passerines)
EPC Costs and Benefits
- both sexes willing
- both leave territories
- maximizes fitness
EPC Benefits in Females
- insurance against infertility
- more genetically diverse brood
- improved genetic quality
- increased access to resources
EPC Costs in Females
- reduced parental care
- increased risk of harassment from mates
- increased risk of STIs
EPC Benefits in Males
- insurance against infertility
- more offspring
- future mate acquisition
EPC Costs in Males
- sperm depletion
- increased risk of divorce/partner infidelity
- reduced parent care
- increased risk of STI
Polygyny
One male, several females, evolved in clumped resource territories
Polygyny Behaviors
- fruit/nectar diets favor polygyny
- heavily associated with lekking
3 Types of Polygyny
1. Female Defense Polygyny
2. Resource Defense Polygyny
3. Male Dominance
resource defense polygyny
males defend territories rich in resources that are used by and attract females
female defense polygyny
Polygynous males directly defend several females.
male dominance polygyny
a mating system in which a few males on a lek mate with many females
male trade-off hypothesis
Polygynous males are so preoccupied with other females they should experience greater levels of cuckoldry (lower paternity rates)
Female choice hypothesis
Females discriminate against solitary males
Comparing phenotypes by the female is easier in groups of males
polyandry
One female, several males (mostly charadriiformes and gruiformes)
hormonal changes in polyandry
sandpiper females have higher testosterone during mating
cooperative polyandry
several males defend a female's territory
brood parasitism
one egg-laying species benefits by having another raise its offspring, eggs develop faster and young fledge sooner
types of brood parasitism
- conspecific/intraspecific (same species)
- interspecific (different species)
conspecific brood parasitism
females facultatively dump their eggs in the nest of other females of their own species
why parasitize conspecifics?
- first step towards evolution of interspecific brood parasitism
- insurance in event female's primary nest is depredated
facultative interspecific brood parasitism
can build own nest, but sometimes dump eggs as well
obligate interspecific brood parasitism
- only way to reproduce; never build own nest (some cowbirds, some cuckoos)
Generalist Brood Parasite
parasitizes multiple different host species
- offspring may kill host eggs
- rejected parasites have nests destroyed
Specialist brood parasites
only parasitizes a single species
defense against brood parasites
- increased nest vigilance
- abandoned nests
cooperative breeding
helpers care for young that is not their own (for experience and inclusive fitness)
Gaining Experience
birds that serve as helpers first have higher lifetime reproductive success
kin selection
the idea that behaviors that help a genetic relative are favored by natural selection
reciprocal altruism
behavior that benefits another with the expectation that those benefits will be returned in the future
why cooperative breeding?
A behavioral pattern in which young birds postpone breeding and instead help their parents raise offspring (in limited locations)
animal architecture
constructions from a variety of natural materials utilizing specific methods to create highly functional designs
nest architecture goals
- protection from predation and parasites
- maintains an adequate microclimate
where do birds build nests?
isolated/hidden places
diversity of nest architecture
- cup
- dome
- dome & tube
- plate/bed
- scrape
- burrow
- mound
nesting materials
many different external materials, sometimes stolen, sometimes none at all
Nest Substrates
- woody vegetation
- grasses
- ground
- wall/ledge
- burrow
- tree cavity
- floating
- animal products
Cavity nesting
- primary (self made)
- secondary (uses other birds')
Weird nesting habits
- spiderwebs by hummingbirds
- hardened saliva by swiftlets
- tree branch by white terns
passerines
most diverse nests, pensile nests
Nesting is inherited
- considered an extended phenotype, but experience matters
evolutionary pressure
nest evolve from avoiding predation
Nest evolutionary history
- began as ground nesting
- diverse in neoaves; adaptive radiation
who builds the nest?
either member, typically both in monogamous species
site selection
important part of pair bonding/mating
nest safety
1. Invisibility
2. Inaccessibility
3. Impregnability
invisibility
- camouflaged eggs
- camouflaged nests
- cryptic locations
inaccessibility
- placed in areas difficult for parasites and predators to get to
impregnability
- physically impenetrable or confusing to enter
nest microclimates
impacted by placement and design