One criticism of natural selection that Darwin anticipated was that it could not easily explain traits that hinder survival (e.g., peacock's tail)
Sexual selection
In The Descent of Man (1871), Darwin elaborated his theory of sexual selection
Sexual selection
Variance in reproductive success results in selection on traits that increase one's ability to win mates
Sexual dimorphism
Caused by the two sexes having different optimal reproductive strategies
Two main processes of sexual selection
Male–male competition for females
Female choice of males
Charles Darwin, 1860: 'The sight of a feather in a peacock's tail, whenever I gaze at it, makes me sick!'
Females
Tend to invest heavily in fewer, larger gametes (eggs)
Males
Tend to invest less in producing many smaller gametes (sperm)
Difference in investment in offspring
Creates conflict in reproductive strategies
Female's fitness
Limited by her own egg/offspring production
Mating with multiple males
Generally provides less fitness benefit for females
Females
Waste valuable eggs if they mate with a low-quality male, so females are expected to be choosy
Male's fitness
Limited by how many eggs he can fertilize
Mating with more females
Can easily increase a male's fitness
Sperm
Cheap to produce, so the cost of mating with a low-quality female is low
On average, making eggs may be ~1000 times more metabolically expensive than making sperm
Variance in reproductive success
Males have much higher variance than females in many species
Females
Generally have about as many offspring as they are physiologically capable of producing
Variation
The fuel for selection, so the sex with the greater variance in mating success will likely experience stronger sexual selection
If the variation is heritable, selection will result in evolution
Strong selection on males
For mating success, may result in evolution of traits for male–male competition or female attraction
Operational sex ratio
When female biased, sexual selection may act on females
Seahorses and pipefish
Females lay eggs into a male's brood pouch, and he provides care until the eggs hatch
Females
Have higher variance in reproductive success and would experience stronger sexual selection
Sexual dimorphism
A difference between the sexes; it ranges from minor to extreme
Intrasexual selection
Competition between members of the same sex (usu. males) for access to members of the opposite sex
Intrasexual selection
Includes physical combat and sperm competition
Physical combat
Male–male competition among ruminants has led to the evolution of headgear that is used as sexual weaponry
Physical combat
Antlers, horns, pronghorns, & ossicones
Physical combat
Sexually dimorphic horns or tusks are also found in other mammals, some insects, fish, reptiles, dinosaurs, etc.
Sperm competition
Competition between males may continue even after copulation
Sperm competition
If a female mates with multiple males, their sperm may compete inside the female by racing to fertilize the egg(s)
Sperm competition
May result in evolution of larger male ejaculations and/or changes to sperm properties (e.g. faster swimming)
Sperm competition
Males may develop penile structures for removing the sperm of other males (e.g., in damselflies)
Males may also deposit a copulatory plug that plugs or glues a female's genital tract closed
Testes size among primates
Most variation is explained by body size, but a significant proportion is explained by mating system
Polygamous species
Have larger testes for their body size than monogamous species, consistent with the hypothesis that larger testes are an adaptation for sperm competition
Alternative mating strategies: the "sneaker male"
When there is intense competition among males over access to mates, alternative "sneaky" mating strategies may evolve
Alternative mating strategies: the "sneaker male"
Smaller, less dominant males may try to obtain access to females while the dominant male is not looking
In some species (e.g., bluegill sunfish, wrasse, cuttlefish), sneakers mimic females to fool the larger male
Infanticide
In some situations, selection may favour killing competitors' progeny
Infanticide
Male lions frequently do this when they immigrate into a pride and take it over