Sexual Selection:

choosy females and

showy males
Mating behavior and mate choice
-Females tend to be choosy, while males
tend to be less choosy
-Females exercise mate choice

-Males compete for opportunities to mate (resources

for breeding)

-Exceptions to this pattern occur, but are rare

Why does this pattern of choosy females and

competitive males exist?
 Natural selection is the differential survival
and reproduction of individuals due to
differences in phenotype

Charles Darwin
 Sexual selection: a form of
natural selection due to
variation in mating success

Charles Darwin

Darwinian Puzzle: elaborate male

behaviors, ornaments, weapons 
Darwin puzzled by existence of traits
that seemed to have no practical
survival advantage

He termed these traits

ornaments

Hypothesized ornaments have

evolved through female choice
 Darwin’s idea: male ornaments evolved
through female choice
females prefer males with the
most extreme ornaments
extreme ornaments =
greater reproductive success

genes for extreme ornaments will

increase in frequency
Sexual selection leads to sexual dimorphism: differences between the
sexes
Darwin: many instances of sexually
dimorphic traits cannot be explained by
ordinary natural selection
These traits might actually make males more
likely to die young

Sexual selection acts on traits that contribute

not to survival, but to an individual’s mating
success

Sexual dimorphism results from

sexual selection acting differently
on males and females
 Two types of sexual selection
Intrasexual selection Intersexual selection
• Within-sex selection • Between-sex selection
• Competition within a sex for mates • Mate choice: preference for particula
• Mainly males individuals of the opposite sex
• Mainly females

Leads to dimorphism due to female

preference for elaborate male traits

Leads to sexual dimorphism

related to weaponry and
competition
 Bateman explained trend toward choosy
females/non-choosy males using relationship
between mating success and reproductive success

A. J. Bateman
Fertility (# progeny)

Reproductive success increased

steeply with number of mates
for males, but not for females

Open squares: male

Closed circles: female

Mating success (# mates)

Bateman gradient (AKA sexual selection gradient):
relationship between reproductive success and mating
success
Bateman’s principle: the sex with the
steeper sexual selection gradient will
Fertility (# progeny)

compete to mate, the other sex will

exercise choice

Open squares: male

Closed circles: female

Mating success (# mates)

 Parental investment hypothesis:
the pattern of mate competition is determined by sex
differences in parental investment
Robert Trivers

Anisogamy:
differences in parental investment
begin with fundamental differences
between male and female gametes
 After eggs are fertilized,
females often spend more
time and energy than
males caring for offspring

Eggs are in limited quantity, large and costly to

produce:
Single egg = 20% of a female’s body mass

In contrast, males produce “cheap” sperm: 8

billion sperm in testes at any given moment Splendid
fairy wren
Male ornaments
evolve through
female choice

females prefer males with the

most extreme ornaments

how do we know that females

care about male ornaments?
Long-tailed widowbirds
Demonstration of female choice for an ornament: long-
tailed widowbird tail length manipulation

Malte Andersson

shortened

cut/glue control
unmanipulated
Males with elongated tails
control recruit more new females to
territory
elongated
Why do females care about tail length?
 Three hypotheses
females, by having preferences,
receive:
Why has
Direct benefits – increase the female’s
female survival, fecundity

preference Indirect benefits – increase fitness of

female’s offspring
evolved? No benefits – evolution of the
preference despite no benefit to the
female
Direct benefits: female obtains something that increases her own
fitness (survival, fecundity)
Alkaloid provides protection from predators

Anti-predation alkaloid

+ = raised with alkaloid

- = raised without alkaloid
 (These are not moths)

Males signal possession of alkaloid

with a pheromone, “HD” Honest signal
of alkaloid possession

This is a spermatophore
Males pass sperm to females in a
spermatophore; contains alkaloid and
nutrients
 70
Number of times male pre- Females benefit from receiving alkaloid
60
ferred by female
50 (gaining protection from predators), and
40 in passing to offspring
30
20
10
0
Yes No
Alkaloid in diet

Females prefer males

Parent raised on PB
with the pheromone diet = no alkaloids

Parent raised on CM
diet = alkaloids
 Three hypotheses: females, by having
preferences, receive:
Why has
female Direct benefits – increase female’s survival,
fecundity

preference Indirect benefits – increase fitness of

female’s offspring via genes from male
evolved? No benefits – evolution despite no benefit to
female
 Preference evolution via indirect benefits:

Sir Ronald Fisher

Good genes Runaway mechanism

• Females prefer a trait • Females prefer some male trait
indicating male’s viability May be arbitrary, trait may not be
Handicap: costly beneficial to female or to offspring
• Selection can “run away” because the
• Benefit: offspring inherit
genes for the female preference and
genes for viability
the male trait become linked
• Benefit: sons inherit genes for
attractiveness: “sexy sons”
Evidence of preference evolving via indirect benefits
Good genes: Advertisement calls of gray tree frogs
Males contribute ONLY sperm
Behavioral tests showed that females prefer longer calls

Experiment: artificially fertilize some eggs w/ long call male sperm, and some eggs
with short call male sperm: compare viability of the offspring

Long call offspring: better performance

Long call males = GOOD GENES
 Three hypotheses, that females, by
having preferences, receive:

Direct benefits – increase female’s

Why has female survival, fecundity
preference evolved? Indirect benefits – increase fitness of
female’s offspring
No benefits – evolution of female
preference despite no benefit to female
or her offspring
Why should a female have a preference if
she gets no benefit from it?
Sensory bias: preference can be result of selection for a different function
of female’s sensory system; males evolve signals to match sensory system
characteristics

Evidence of preference evolving via no benefits:

sensory bias in swordtails

-No evidence for use of sword in male-male

competition
-Not likely to be useful in swimming
-Not helpful in avoiding predation
-Likely evolved through female choice
Tested preference for swords in a species where
males have swords, X. helleri

Male Male
Female

Females prefer longer swords

Prediction: if preference for swords is due to a

sensory bias, the preference for swords must have
evolved before the sword trait itself
To show that the
preference evolved
before the sword trait,
must demonstrate
preference in genus
Priapella
Surgically implanted swords into a
preference species without swords, Priapella

Priapella females prefer

Time spent near male (sec)

Possible males w/ swords!
origin of
trait
Possible
origin of
trait
Sexual Selection:
Showy and competitive males
 Intrasexual selection:
within-sex competition
Typically males fight with males:
-Larger male body size (sexual dimorphism)
-Armaments (weapons – sexual dimorphism)
-Male dominance hierarchy: a few males are most
likely to mate

Consider cost of achieving high status vs.

reward of fathering more offspring
Reproductive benefit must outweigh
the cost of competition
 In elephant seals, mating success is highly
correlated with dominance rank

• Males are sexually mature 5-6 yrs

• Do not achieve alpha status until 9 yrs
• Alpha status 9-12 yrs
• Typical lifespan 14 yrs

Early years spent GROWING, learning to fight

Most will never achieve alpha status
Alpha status is short-lived
Die young

The costs of this life-history strategy are high!

What are the benefits?
 In elephant seals, mating
success is highly correlated with
dominance rank

Most dominant males get

most of the reproductive
success
The relationship between dominance and sexual access: baboons

For most baboon

groups: very high
correlation between
dominance rank and
mating success

Hierarchy is important in baboons

Cost: Males get bitten every 6 weeks (4x female injury rate)
But….. high ranking males dominate copulations when females are
most fertile
intrasexual selection: sperm competition
competition between males that determines whose sperm will fertilize a
female’s eggs

Male damselflies remove sperm of previous

partners before passing their sperm to female
In some species females can affect which Female collared flycatcher
sperm fertilizes her eggs
Females may mate with males other
than their social partner by seeking
extra-pair copulations

Sperm can be stored

up to a month

By controlling when she

mates with whom, females
can bias fertilization toward
particular males

A female paired with a male with a small

white forehead patch can secure sperm
around the time of egg laying by slipping
away to mate with a nearby male who sports
Male collared flycatcher a larger white patch