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Chapter 10
Mating Systems

10.1 Monogamy: A Lack of Multiple Mating

10.2 Polyandry: Multiple Mating by Females
10.3 Polygyny: Multiple Mating by Males
10.4 Polygynandry and Promiscuity: Multiple Mating by Both Sexes

10.1 Monogamy: A Lack of Multiple Mating

LEARNING OBJECTIVES
■ Review the costs and benefits of mating monogamously for males and females across different taxa.
■ Compare the various hypotheses for the evolution of monogamy, emphasizing the inability for either sex to
monopolize access to mates or resources.
■ Describe the relationships among biparental care, mate guarding and mate assistance, and the evolution of social
monogamy.

Q: What does social monogamy in birds typically imply about genetic monogamy?
A: Social monogamy in birds does not necessarily equate to genetic monogamy. In the vast majority of bird species,
one or both members of a pair-bond engage in extra-pair fertilizations, meaning they produce offspring with
individuals outside their social pair-bond. Therefore, while birds may form social monogamous pair-bonds, they often
exhibit genetic polygamy, with extra-pair fertilizations occurring frequently.

Q: What did Charles Nunn and his team investigate regarding mating systems in primates?
A: Charles Nunn and his team investigated whether there's a relationship between mating systems and immune
system strength in primates.

Q: How did they conduct their study?

A: They used a comparative approach, comparing species with varying mating systems, ranging from extremely
polyandrous to monogamous.

Q: What were some of the costs associated with mating multiply?

A: Costs of mating multiply can include the time and energy spent searching for and mating with several partners, the
risk of being killed by a predator during these forays and when mating, and the chance of acquiring a sexually
transmitted disease from some mates.

Q: What prediction did they make about the immune systems of species that mate multiply?
A: They predicted that the immune systems of species that mate multiply would be stronger than those of related
species that have a greater tendency toward monogamy.

Q: What did they find in their study regarding the relationship between mating systems and immune system strength?
A: They found that in species with more polyandrous mating systems, such as the Barbary macaque, females mate
with as many as ten males in one day, and their white blood cell counts were indeed higher (but within the normal
range) compared to monogamous species.
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Q: What factors contribute to the costs of mating multiply?

A: The costs of mating multiply can include:

Time and energy spent searching for and mating with multiple partners.
Increased risk of predation during forays and mating.
Higher likelihood of acquiring sexually transmitted diseases from multiple mates.

Q: How did Nunn and his colleagues measure immune system strength across different mating systems in primates?
A: They measured immune system strength by analyzing white blood cell counts from adult females of various primate
species, most of which were monitored for health in zoos.

Q: What did they find regarding the relationship between mating systems and immune system strength in primates?
A: They found that in species with more polyandrous mating systems, the white blood cell counts of healthy females
were higher, suggesting a potential link between mating multiply and immune system strength.

Q: Was this relationship specific to primates, or did similar patterns emerge in other animal groups?
A: Similar patterns were found in other animal groups. In carnivores, there was also a relationship between white blood
cell count and the number of mating partners a female had. Additionally, in birds, polyandrous species had higher
levels of sexually transmitted bacteria and yeast compared to monogamous species.

FIGURE 10.2 Polyandry has fitness costs. In primates, the number of mates accepted by females is correlated with the
investment made in white blood cells, a component of an animal’s immune system. (The higher the measure of white
blood cells, the stronger the immune system.) This result suggests that the more polyandrous the species, the greater
the challenge to the female’s immune system. Each data point represents a different primate species. Only species
with well-defined periods of female mating activity were included in the analysis. Negative values are the result of
controlling for other variables and for evolutionary history in the analysis. (After C. L. Nunn et al. 2000. Science 290:
1168–1170.)

Q: What are some hypotheses proposed to explain the evolution of monogamy?

A: The hypotheses proposed to explain the evolution of monogamy include the mate limitation hypothesis, mate
guarding hypothesis, mate assistance hypothesis, and infanticide hypothesis.

Q: Can you explain the mate limitation hypothesis?

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A: The mate limitation hypothesis suggests that monogamy evolves when potential mates are not readily available in
groups and instead roam widely, making them costly to locate.

Q: What is the mate guarding hypothesis?

A: The mate guarding hypothesis proposes that monogamy evolves when individuals have the ability to restrict mating
behavior in their partner, thus ensuring fidelity and reducing the risk of losing reproductive opportunities.

Q: Could you elaborate on the mate assistance hypothesis?

A: The mate assistance hypothesis suggests that monogamy evolves when resources are crucial for successful
reproduction, requiring both parents to contribute to the rearing of offspring.

Q: What does the infanticide hypothesis propose?

A: The infanticide hypothesis suggests that monogamy evolves when the risk of infanticide is high, and having a
partner can provide protection against infanticidal males.

Q: What are some non-mutually exclusive hypotheses to explain the evolution of monogamous mating systems?
A: The non-mutually exclusive hypotheses include the mate limitation hypothesis, mate guarding hypothesis, mate
assistance hypothesis, and infanticide hypothesis.

Q: What does the mate limitation hypothesis propose?

A: The mate limitation hypothesis suggests that monogamy is likely to evolve when potential mates do not form
groups and roam widely, making them costly to locate.

Q: Can you explain the mate guarding hypothesis?

A: The mate guarding hypothesis suggests that monogamy is likely to evolve when individuals have the ability to
restrict mating behavior in their partner, thus ensuring fidelity and reducing the risk of losing reproductive
opportunities.

Q: What is the mate assistance hypothesis?

A: The mate assistance hypothesis proposes that monogamy is likely to evolve when resources are crucial for
successful reproduction, requiring both parents to contribute to the rearing of offspring.

Q: Could you elaborate on the infanticide hypothesis?

A: The infanticide hypothesis suggests that the risk of infanticide by males may influence female reproductive tactics
and the evolution of monogamy by forming a bond with a partner to provide protection for offspring.

Q: How does mate guarding behavior relate to the mate guarding hypothesis?
A: Mate guarding behavior, where males prevent their mates from seeking extra-pair fertilizations, aligns with the
mate guarding hypothesis, which suggests that monogamy evolves when individuals have the ability to restrict their
partner's mating behavior to ensure fidelity.

Q: What is an example of a species where males exhibit mate guarding behavior?

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A: Some species of spiders exhibit mate guarding behavior, where males give their all to their first mate, even
engaging in sexual suicide by breaking off their genital appendages or offering themselves as food to their partner, to
increase their fertilization success with that one female.

Q: How does parental care contribute to the mate assistance hypothesis?

A: Parental care plays a central role in the mate assistance hypothesis, which proposes that a male remains with a
single female to help her with various tasks, particularly because paternal care and protection of offspring are
advantageous.

Q: Can you describe an example of paternal care contributing to the mate assistance hypothesis?
A: In a study of crickets, males were observed to guard their mates, significantly reducing the likelihood of the females
falling victim to predators. Despite increasing his own risk of being killed by allowing the female to dash into the
burrow first, the male ensured his paternity by mating repeatedly with the female, thus increasing the survival odds of
the female carrying his offspring.

FIGURE 10.3 Mate guarding and mate assistance in the cricket Gryllus campestris. (A) A male that has acquired a mate
remains with her in his territory. If the pair is attacked, the male permits the female to rush into the burrow first, at
great cost to his personal safety. (B) Solitary territorial males are much more likely to survive an attack. A female that
lives alone is less likely to survive than one paired with a self-sacrificing partner. (B after R. Rodríguez-Muñoz et al.
2011. Curr Biol 21: 1716–1719.)

Q: What is female-enforced monogamy?

A: Female-enforced monogamy refers to a form of mate guarding where a female prevents her partner from engaging
in polygynous behavior, thereby ensuring his exclusive parental assistance.

Q: Can you describe an example of female-enforced monogamy in burying beetles?

A: In the burying beetle species Nicrophorus defodiens, paired females are aggressive toward intruding females. If a
mated male releases a sex pheromone to attract a second female to the site where they buried a dead mouse or
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shrew, the paired female may push him from his perch to prevent another female from adding her clutch of eggs to
the carcass. This behavior reduces the male's ability to signal and may result in monogamy enforced by females.

Q: How did Anne-Katrin Eggert and Scott Sakaluk demonstrate the impact of female mate guarding on male signaling
behavior in burying beetles?
A: Anne-Katrin Eggert and Scott Sakaluk conducted an experiment where they tethered paired females of burying
beetles to prevent them from suppressing their partners' sexual signaling. They found that males whose partners were
not able to control them released scent for longer periods compared to control males with untethered mates,
demonstrating the influence of female mate guarding on male signaling behavior.

FIGURE 10.4 Female-enforced monogamy in burying beetles. (A) When a paired female beetle (B) is experimentally
tethered so that she cannot interact with her partner, the amount of time he spends releasing sex pheromones rises
dramatically. (After A.-K. Eggert and S. K. Sakaluk. 1995. Behav Ecol Sociobiol 37: 147–154.)

Q: What is the relationship between monogamy and biparental care in bird species?
A: In many bird species, males and females form partnerships for one or more breeding seasons, and males contribute
significantly to the welfare of the offspring produced by their mates. This biparental care is essential for the survival of
the young and is often associated with social monogamy.

Q: Can you provide an example of paternal assistance in bird species?

A: In the yellow-eyed junco (Junco phaeonotus), the male takes care of his mate’s first brood of fledglings while the
female incubates a second clutch of eggs. This paternal help is crucial for the survival of the fledglings, which are
highly inept foragers.

Q: How does biparental care impact the development of offspring in European starlings?
A: In European starlings (Sturnus vulgaris), clutches with biparental attention stay warmer and develop more rapidly
compared to clutches cared for by mothers alone. Research has shown that 97 percent of the eggs incubated by both
parents hatch, compared to only 75 percent of eggs cared for by mothers alone.
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FIGURE 10.5 Paternal male European starlings help their mates. Males keep their clutches warmer by sharing in
incubation duties. (A) Eggs that were incubated by both parents were kept at about 35°C most of the time, whereas
eggs incubated by the female alone were often several degrees cooler. (B) Both male and female starlings incubate
the eggs. In this nest, all the eggs hatched successfully. (A after J. M. Reid et al. 2002. Behav Ecol Sociobiol 51: 255–
261.)

Q: What is the impact of experimental "widowing" on the reproductive success of snow buntings?
A: Studies on widowed snow buntings (Plectrophenax nivalis) have shown that females left to rear broods on their own
usually produce three or fewer young, whereas control pairs often fledge four or more, indicating that reproductive
success is closely linked to paternal care behavior in many monogamous species.

Q: How does the administration of extra testosterone affect paternal care behavior in male spotless starlings?
A: Male spotless starlings (Sturnus unicolor) given extra testosterone become less willing to feed nestlings, whereas
males administered an anti-androgenic chemical that blocks the effects of naturally circulating testosterone feed their
offspring at increased rates.

Q: What is the relationship between the administration of extra testosterone and the number of fledged young per
brood in spotless starlings?
A: The mean number of fledged young per brood was lowest for spotless starlings with extra testosterone and highest
for those administered the testosterone blocker, indicating that testosterone levels can influence paternal care
behavior and ultimately affect reproductive success.
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FIGURE 10.6 Paternal care boosts reproductive success in spotless starlings. (A) Male spotless starlings whose (B)
testosterone levels were reduced by the anti-androgen cyproterone acetate (CA) provided more food for their broods
and had the highest fledging rates per brood. Males given extra testosterone (T) provided less food and had the lowest
fledging rates. Untreated controls were intermediate with respect to both feeding and fledging rates. Values depict
mean ± SE. (After J. Moreno et al. 1999. Behav Ecol Sociobiol 47: 47–53.)

Q: What unique parental behavior is exhibited by male seahorses such as Hippocampus whitei?
A: Male seahorses, like those of the species Hippocampus whitei, take on the responsibility of "pregnancy" by carrying
a clutch of eggs in a sealed brood pouch for about 3 weeks.

Q: Describe the social behavior of male and female seahorses in terms of their mating habits and interactions.
A: Male and female seahorses, such as those in the species Hippocampus whitei, have durable relationships, with each
male typically paired with a single female that provides him with a series of clutches. Pairs of seahorses even greet
each other each morning before moving apart to forage separately, and they ignore any members of the opposite sex
they encounter during the day.

Q: How does the reproductive behavior of male seahorses such as Hippocampus whitei limit their potential mates?
A: The reproductive behavior of male seahorses limits their potential mates because a male's brood pouch can
accommodate only one clutch of eggs, meaning he gains nothing by courting more than one female at a time. In some
species, like another type of Hippocampus, genetic data indicate that males do not accept eggs from more than one
female, even if females attempt to court them.
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FIGURE 10.7 Monogamy and paternal care in a seahorse. A pregnant male is giving birth to his single partner’s
offspring. Two young can be seen emerging from their father’s brood pouch.

Q: How does the reproductive behavior of male seahorses align with the concept of monogamy?
A: Male seahorses may not benefit from switching mates if their long-term mate can supply them with a new
complement of eggs as soon as one pregnancy is over. By pairing off with one female, a male seahorse can match his
reproductive cycle with that of his partner, allowing him to immediately secure a new clutch from his familiar mate.

Q: How do experiments with paternal pipefish support the idea of monogamy in relation to reproductive intervals?
A: Experimental removal of a female partner in paternal pipefish lengthened the interval between spawning for the
male. Males that had to change mates took significantly longer to acquire replacement partners with clutches of
mature eggs compared to males permitted to retain their mates from one clutch to the next.

Q: Why is monogamy exceptionally rare in mammals, according to sexual selection theory?

A: Monogamy is exceptionally rare in mammals because sexual selection theory suggests that males should try to
mate with many females due to the limited types of care a male can provide offspring compared to a female,
particularly at the earliest developmental stages.

Q: How do exceptions to the rarity of monogamy in mammals, such as the Djungarian hamster and the California
mouse, contribute to understanding the mate assistance hypothesis?
A: Exceptions to the rarity of monogamy in mammals, such as the Djungarian hamster and the California mouse,
support the mate assistance hypothesis for monogamy. Males of these species exhibit paternal behavior and
contribute to offspring survival, suggesting that male monogamy may be associated with providing assistance in
rearing offspring.
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FIGURE 10.8 An exceptionally paternal mammal, the Djungarian hamster. A male Djungarian hamster may pull
newborns from his mate’s birth canal and then clear the infants’ airways by cleaning their nostrils, as shown in these
photographs (the male is the hamster on the left; arrows point to the pink newborn).

FIGURE 10.9 Male care of offspring affects fitness in the California mouse. Although the number offspring born to
mothers in each treatment group did not differ, the number of offspring reared until first emergence from the nest by
female mice falls sharply in the absence of a helpful male partner. Bars depict mean ± SE. (After D. J. Gubernick and T.
Teferi. 2000. Proc R Soc London B 267: 147–150.)

Q: How does the infanticide hypothesis explain the evolution of monogamy in mammals?
A: According to the infanticide hypothesis, monogamy may be favored when the risk of infanticide is high because
having a partner can help provide protection against infanticidal males. Paternal males in some mammal species may
drive away other males from their brood, thereby blocking potential infant killers and preventing infanticide.
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Q: How does the mate-guarding prairie vole demonstrate the contributions of both mate guarding and mate assistance
to monogamy?
A: In the mate-guarding prairie vole, males can and do defend their offspring against infanticidal rivals, demonstrating
the role of mate guarding in monogamy. Additionally, paternal care behaviors contribute to the tendency toward
monogamy in this species.

Q: How does infanticide risk influence monogamy in primate species?

A: Infanticide risk may favor monogamy in many primate species. For example, in the white-handed gibbon, groups
composed of a monogamous pair living with their offspring have a much lower risk of infant disappearance compared
to polyandrous groups consisting of an adult female and several adult males.

Q: What did Dieter Lukas and Tim Clutton-Brock find when testing the infanticide hypothesis in mammals?
A: Lukas and Clutton-Brock found no relationship between monogamy and the risk of male infanticide in more than
2500 species of mammals.

Q: How did Opie and colleagues respond to Lukas and Clutton-Brock's findings regarding the relationship between
male infanticide risk and monogamy in mammals?
A: Opie and colleagues countered Lukas and Clutton-Brock's findings by suggesting that the lack of evidence for the
relationship between male infanticide risk and monogamy in mammals might be due to taxonomic differences. They
argue that infanticide risk in primates is driven by their unusually late weaning, unlike males of other mammal species
with shorter development periods, making infanticide an unlikely selection factor for monogamy outside of primates.

Q: What is Lukas and Clutton-Brock's perspective on the relationship between paternal care and monogamy in
mammals?
A: Lukas and Clutton-Brock argue that paternal care is a consequence rather than a cause of monogamy in mammals.
They suggest that monogamy has evolved in mammalian species where breeding females are intolerant of each other
and where female density is low, consistent with the mate guarding hypothesis.
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FIGURE 10.10 Female density predicts monogamy in mammals. Although the values for solitary species (blue dots)
and monogamous species (red squares) overlap, monogamy tends to occur in mammalian species where female
density is low (logarithm of the number of individuals per km2). At the highest population densities, there is only a 6
percent probability that a species will be monogamous, whereas the probability rises to 44 percent at the lowest
population densities. The dashed line indicates the fit from a statistical model controlling for other life history traits.
(After D. Lukas and T. H. Clutton-Brock. 2013. Science 341: 526–530.)

Self-Study Questions 10.1

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1. Although monogamy is extremely rare, its primary benefits are

decreased chances of inbreeding.

paternity assurance and increased survival of offspring.

increased access to mates for males.

increased odds of receiving compatible genes.

more heterozygous offspring and higher offspring immunity.

2. In a study on monogamy in mammalian species, Lukas and Clutton-Brock found that monogamy is more likely to
evolve when female density is low. This finding best supports the

inbreeding avoidance hypothesis: females choose mates to avoid inbreeding and produce more heterozygous
offspring.

mate guarding hypothesis: males are unable to gain and defend access to multiple females.

mate assistance hypothesis: both parents are necessary to provide parental care for young.

infanticide hypothesis: males are needed to provide protection against infanticidal rivals.

good genes hypothesis: females receive genetic benefits of improved offspring quality and viability.

3. Under both the mate assistance hypothesis and the infanticide hypothesis, monogamy is likely to evolve when

individuals have the ability to restrict mating behavior in their partner.

it is costly to locate potential mates because they roam widely and do not form groups.

there is little male-male competition.

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paternal care and protection increase survival of offspring.

there is little chance for female extra-pair copulations.

10.2 Polyandry: Multiple Mating by Females

LEARNING OBJECTIVES
■ Differentiate the degree of control over mating between males and females in polyandrous systems, considering the
reproductive costs and benefits, as well as the ecological factors, driving the evolution of polyandry.
■ Review the potential indirect genetic benefits gained by females for their offspring by mating polyandrously.
■ Evaluate the hypotheses for the potential direct benefits that females obtain by mating with multiple males.

Q: What are some potential benefits to females of being choosy in selecting mates?
A: Potential benefits to females of being choosy include indirect genetic advantages to their offspring and direct
benefits such as parental care, access to resources, safety from predators, or reduced harassment by other males, all
of which can increase a female’s fitness.

Q: What surprised ornithologists regarding the fidelity of female birds and why?
A: Ornithologists were surprised to find that many female birds are not faithful because they had assumed that males
helped their mates care for their mutual offspring, not for the progeny of other males.

Q: What did a review led by Nina Wedell find regarding mating with more than one male per female across various
taxonomic groups?
A: The review found evidence that mating with more than one male per female occurs in all of the 14 major taxonomic
groups explored, from sea spiders to mammals, and in 89 percent of populations examined.

Q: What did researchers discover in a study of pied flycatchers regarding female flying ability and extra-pair
fertilizations?
A: Researchers found that experimentally decreasing female flying ability resulted in nearly double the proportion of
extra-pair young in their nests compared to control females. These offspring were sired by more males than those in
control broods that contained extra-pair young, suggesting that females exert control over with whom they mate.

Q: How does the existence of constraints imposed by choosy females and competitors help explain the evolution of
polyandrous mating systems?
A: The existence of constraints imposed by choosy females and competitors may explain the evolution of polyandrous
mating systems where most breeding females form social bonds with several males. In species like the Galápagos
hawk, polyandry is associated with a scarcity of suitable territories, leading to a highly male-biased operational sex
ratio. This intense competition for females has favored males capable of forming cooperative defense teams, resulting
in male harems that help females rear offspring.

Q: What is a notable feature of the wattled jacana regarding its mating system?
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A: In the wattled jacana, a polyandrous species, females aggressively fight for and defend territories that can
accommodate several males. The territorial female mates with all members of her harem, providing each male with his
own clutch of eggs, which he cares for exclusively.

Q: What happens in the red phalarope regarding parental care and offspring paternity?
A: In the red phalarope, a polyandrous shorebird, females fight for males, and a male provides all parental care for the
clutch deposited in his nest, regardless of whether the eggs have been fertilized by other males. This results in males
caring for offspring sired by other males, illustrating a cost to males that are pair-bonded with a polyandrous female.

FIGURE 10.11 Female red phalaropes are polyandrous. After securing one male partner, a female phalarope may
attract another, donating a clutch of eggs to each male in turn. The more brightly colored bird (on the left) is the
female.

Q: What ecological factors contribute to the mating system of the spotted sandpiper?
A: The mating system of the spotted sandpiper is influenced by several ecological features. Firstly, the adult sex ratio
is slightly biased toward males, limiting opportunities for polygyny and prompting females to compete for mates.
Secondly, these birds nest in areas with abundant food from mayfly hatches, allowing for multiple clutches to be laid.
Thirdly, because spotted sandpiper chicks are precocial, a single parent can care for a clutch as effectively as two
parents.

Q: How do female spotted sandpipers affect their mates in terms of reproduction?

A: Female spotted sandpipers may desert their initial partners, but this does not harm the survival chances of their
first broods due to the abundance of food and the precocial nature of the young. Once deserted, a male is left stuck,
as leaving the nest would cause the eggs to fail to develop. However, if all females desert, a male single parent may
still experience greater reproductive success. Additionally, a female may store sperm from the first male to mate with
her and use it later to fertilize her second clutch of eggs.

FIGURE 10.12 Polyandrous female spotted sandpipers fight for males. Two female spotted sandpipers (A) about to fight
and (B) fighting for a territory that may attract several monogamous, paternal males to the winner.

Q: How do male red-necked phalaropes maintain some control over their mating decisions in polyandrous systems?
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A: Male red-necked phalaropes favor their original sexual partners over novel females, reducing the risk of caring for
eggs fertilized by other males. When polyandrous females draw second mates from males that have lost a first clutch
to predators, the male still prefers his original partner over a new female.

Q: How do male red jungle fowl strategically allocate sperm in polyandrous mating scenarios?
A: In polyandrous red jungle fowl, dominant males increase their sperm investment as the number of mates a female
has increases, to reduce the risk of another male inseminating the same female. Subdominant males, on the other
hand, maximize their sperm investment when faced with just one competitor, potentially saving sperm for future, less
competitive copulation opportunities.

FIGURE 10.13 Male red jungle fowl adjust their sperm investment according to how many other mates a female has.
Relative sperm investment of dominant (dark blue bars) and subdominant (light blue bars) males according to female
promiscuity: dominant males steadily increased relative sperm investment as the number of competitors increased,
whereas subdominants decreased their investment when the number of competitors increased beyond one. Relative
sperm investment was calculated as the cumulative number of sperm transferred by a male during a trial standardized
against his largest cumulative number of sperm produced in a trial across all audience treatments. Bars depict mean ±
SE. (After T. Pizzari et al. 2003. Nature 426: 70–74.)

Q: How do males in polyandrous mating systems maintain some control over mating despite females typically exerting
more control?
A: Males in polyandrous mating systems have evolved mechanisms to maintain some control over mating. For
example, in chinook salmon, subordinate males produce seminal fluid that increases the swimming speed of sperm
from dominant males, while the seminal fluid from dominant males decreases the swimming speed of sperm from
subordinate males. This difference in sperm velocity affects the proportion of eggs sired, with sperm incubated in
subordinate male seminal fluid siring a greater proportion of eggs.
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FIGURE 10.14 Seminal fluid of the chinook salmon influences sperm swimming speed. The average difference in sperm
velocity (VAP) between sperm incubated in their own seminal fluid and those incubated in the seminal fluid of a rival.
Seminal fluid from dominant rival males decreased the velocity of sperm from subordinate males, whereas seminal
fluid from rival subordinate males increased the velocity of sperm from dominant males. Bars depict mean ± SE. (After
M. J. Bartlett et al. 2017. eLife 6: e28811. CC BY 4.0.)

Q: What are some potential benefits for females from mating multiply?
A: Females can potentially obtain several indirect genetic benefits for their offspring by mating polyandrously and
obtaining sperm (and genes) from multiple males.

Q: What is the good genes hypothesis?

A: The good genes hypothesis suggests that females mate polyandrously to produce offspring of higher genetic quality
or viability.

Q: What is the genetic compatibility hypothesis?

A: The genetic compatibility hypothesis proposes that females mate polyandrously to increase the odds of receiving
genetically complementary sperm.

Q: What is the genetic diversity hypothesis?

A: The genetic diversity hypothesis suggests that females mate polyandrously to increase the heterozygosity (genetic
diversity) of either individual offspring or of the group of offspring produced in a single bout.

Q: What is the inbreeding avoidance hypothesis?

A: The inbreeding avoidance hypothesis posits that females mate polyandrously to avoid inbreeding with their social
partner.

Q: What is the main idea behind the good genes hypothesis?

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A: The main idea behind the good genes hypothesis is that females mate polyandrously to produce offspring of higher
genetic quality or viability.

Q: Can you provide an example that supports the good genes hypothesis?
A: Yes, an example supporting the good genes hypothesis is the grey foam nest treefrog (Chiromantis xerampelina),
where offspring from the nests of polyandrous females were significantly more likely to survive to metamorphosis than
those of females with just a single mate.

FIGURE 10.15 Polyandry has fitness benefits. (A) Females of the grey foam nest treefrog lay clutches of eggs that may
be fertilized by anywhere from 1 to 12 or more males. The offspring of polyandrous females are more likely to survive
than are those of females mated to only one male. Notice the large number of males trying to fertilize a single
female’s eggs that have already been laid inside the foam nest. (B) A male yellow-toothed cavy, whose very large
testes speak to the intense sperm competition in this species, which occurs because females usually mate with several
males. (C) When female yellow-toothed cavies were experimentally restricted to a single mate, they had fewer
surviving offspring than females allowed to mate freely with four males. View a larger version of this figure (C after A.
Keil and N. Sachser. 1998. Ethology 104: 897–903.)
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Q: What additional benefits besides improved early survival can polyandry provide for the offspring of females?
A: Polyandry can provide benefits such as increased lifetime reproductive success for offspring, particularly in species
like the dark-eyed junco, where the genetic offspring of females and their extra-pair mates are more likely to survive
to reproduce. Additionally, the offspring from extra-pair liaisons may be more fecund and have higher reproductive
success. These benefits are attributed to the genes contributed by extra-pair males.

Q: What is an alternative explanation for the observed effects of polyandry in species like the dark-eyed junco?
A: An alternative explanation is that when a female mates with an extra-pair male, she may subsequently invest more
in the offspring of such a relationship. This investment can include producing larger eggs fertilized by the sperm of
extra-pair mates, leading to an early developmental boost for the offspring and resulting in fitter offspring. These
effects are attributed to maternal effects rather than acquiring good genes from males.

Q: How did Magdalena Kozielska and her team design their experiment to investigate the potential roles of good genes
versus maternal effects in the bulb mite Rhizoglyphus robini?

A: Kozielska and her team eliminated the possibility of genetic benefits of polyandry by allowing only one randomly
chosen male to sire the offspring of polyandrous females, achieved by sterilizing the other males with ionizing
radiation. They compared the fecundity of daughters of mothers mated monogamously to those mated pseudo-
polyandrously, finding no difference, indicating that previous results were due to the genetic benefits of mating with
multiple males and not to maternal effects. Interestingly, they found maternal effect benefits for sons, as the
reproductive success of sons of pseudo-polyandrous mothers was less than that of sons of monogamous mothers,
even though both sets of males shared the same fathers.

Q: What is the "sexy sons hypothesis" proposed by Weatherhead and Robertson (1979)?
A: The "sexy sons hypothesis" suggests that one benefit females gain from engaging in extra-pair copulations is the
production of sons who inherit traits that make them sexually appealing, thus increasing their reproductive success.

Q: What evidence supports the "sexy sons hypothesis"?

A: Research in the field cricket Gryllus bimaculatus demonstrated that sons of sexually successful males were more
appealing to females compared to sons of males that failed to acquire a mate in experimental settings (Wedell and
Tregenza, 1999). This suggests that male attractiveness can be inherited, supporting the hypothesis.

FIGURE 10.16 A father’s mating success can be transmitted to his sons. In experiment 1, two male field crickets were
given an opportunity to compete for a female; one male (S) mated successfully, while the other male (U) was
unsuccessful. When the sons of male S were placed in competition for a female with the sons of male U (which had
been given a female to mate with after failing to win the initial competition), the sons of S were nearly twice as likely
to mate with the female as were the sons of U. In experiment 2, a male that had won a mating competition was later
allocated a female at random for breeding, as was a male that had lost the competition. The sons of the two males
were then placed in an arena with a female, and as before, the sons of S were much more likely to mate with the
female than were the sons of U. (After N. Wedell and T. Tregenza. 1999. Evolution 53: 620–625.)
18

Q: How did Gwinner and Schwabl investigate the "sexy sons hypothesis" in European starlings?
A: They studied sons of monogamous and polygynous males, examining their nest-site-acquisition behavior and
hormone levels in eggs to rule out potential maternal effects.

Q: What were the findings of the study regarding the behavior and hormone levels of the sons?
A: Sons of polygynous fathers defended more nest boxes and produced more courtship song compared to sons of
monogamous fathers. However, hormone levels in eggs fathered by polygynous versus monogamous males did not
differ, suggesting that these behavioral qualities are inherited from the fathers rather than being influenced by
maternal effects from hormones in the eggs.

Q: What is the genetic compatibility hypothesis regarding polyandry?

A: The genetic compatibility hypothesis suggests that females engage in polyandry to increase the likelihood of
receiving genetically complementary sperm, which can result in highly viable offspring.

Q: How was the genetic compatibility hypothesis tested in pseudoscorpions?

A: The hypothesis was tested by comparing the number of surviving offspring produced by polyandrous
pseudoscorpions with those produced by females paired with lone males. The study found that the reproductive
success of females depended on the match between their gametic genomes and the sperm they received from males.
There was no correlation between a male's reproductive success and the number of offspring produced by different
females from matings with the same male.

Q: What did Jeanne Zeh find when comparing the reproductive success of female pseudoscorpions with different
males?
A: Jeanne Zeh found no correlation between the numbers of offspring produced by different females from matings with
the same male, indicating that male pseudoscorpions cannot be categorized into studs and duds. Instead, the
reproductive success of females depended on the compatibility between the gametic genomes of the male and
female, supporting the genetic compatibility hypothesis.
19

Q: How did female pseudoscorpions respond when given an opportunity to mate with the same male again after 90
minutes?
A: When given the opportunity to mate with the same male again after 90 minutes, female pseudoscorpions refused to
accept his spermatophore in 85 percent of the trials. Instead, they usually accepted the gametes of a new partner,
suggesting a preference for new mates to increase the chances of securing genetically compatible sperm.

FIGURE 10.17 Polyandry boosts female reproductive success in a pseudoscorpion. In laboratory experiments, female
pseudoscorpions restricted to lone partners produced fewer nymphs than females that mated with several males.
Paternity tests of the offspring of wild-caught females confirmed that females usually mate with several males under
natural conditions. Bars depict mean ± SE. (After J. A. Zeh. 1997. Behav Ecol Sociobiol 40: 111–118.)

Q: What is the genetic diversity hypothesis?

A: The genetic diversity hypothesis suggests that females benefit by increasing the heterozygosity (genetic diversity)
of their offspring. This can occur at the individual level, where offspring with two different forms of a gene often have
an advantage over homozygotes, and at the group level, where a greater diversity of individuals can help withstand
disease outbreaks or survive in various environmental conditions.

Q: How might genetically similar pairs be affected by the genetic diversity hypothesis?
A: Genetically similar pairs may be at risk of producing inbred offspring with genetic defects. Therefore, females
bonded with genetically similar individuals may prefer to mate with males who are either genetically dissimilar or
highly heterozygous themselves. This preference increases the likelihood of producing offspring with increased
heterozygosity, which can provide advantages in terms of genetic fitness and adaptability.

Q: How has the genetic diversity hypothesis been tested in bird species?
A: The genetic diversity hypothesis has been tested in bird species such as superb starlings and bluethroats. In superb
starlings, females seeking extra-pair mates outside their social groups tended to have less heterozygous social
20

partners, while offspring fathered by extra-pair males were more heterozygous than those from social partners.
Similarly, in bluethroats, extra-pair offspring were more heterozygous than those sired by social partners.

Q: What additional trait was examined in bluethroats to assess the impact of offspring heterozygosity?
A: In bluethroats, the authors examined whether offspring heterozygosity enhanced immune function. They found that
extra-pair offspring had stronger immune systems compared to offspring sired by social partners. This was
demonstrated by measuring the swelling at the injection site after injecting a foreign substance that triggers an
immune response into the wing webs of nestling bluethroats.

FIGURE 10.18 The effect of extra-pair matings on the genetic heterozygosity of offspring of superb starlings. In nests
containing extra-pair young sired by males from outside the social group (extra-group extra-pair young), the offspring
sired by the extra-pair mate (dark blue bars) had higher heterozygosity than those sired by the social partner (light
blue bars). However, in nests containing extra-pair young sired by males from within the social group (within-group
extra-pair young), there was no difference in heterozygosity between offspring sired by the extra-pair mates and social
partners. Bars depict mean ± SE. (After D. R. Rubenstein. 2007. Proc R Soc B 273: 1895–1903.)

FIGURE 10.19 Extra-pair matings can boost the immune responses of offspring in the bluethroat. The mean wing-
swelling response (an indicator of immune system strength) of young sired by their mother’s social partner (within-pair
young, or WPY) was less than that of young birds sired by their mother’s extra-pair partner (extra-pair young, or EPY)
in broods of mixed paternity. Bars depict mean ± SE. View a larger version of this figure (After A. Johnsen et al. 2000.
Nature 406: 296–299.)
21

Q: How is polyandry observed in some species of the genus Apis, such as A. florea and A. dorsata?
A: In species like A. florea and A. dorsata, females are highly polyandrous. For instance, A. florea females use sperm
from about 10 males on average, while A. dorsata females acquire sperm from an average of 63 males.

Q: What distinguishes the mating behavior of queens in the genus Apis from other bee species?
A: In most bee species, queens typically mate just once. However, in the genus Apis, including species like A. florea
and A. dorsata, virgin queens engage in multiple matings during nuptial flights, being "captured" and mated by several
males in quick succession.

Q: What are some potential individual-level and colony-level benefits proposed to explain the replacement of
monogamy by extreme polyandry in some bee species?
A: Individual-level benefits include long-lived queens not running out of sperm, while colony-level benefits suggest that
genetic diversity among individuals within a colony is more crucial than diversity within any single individual. These
colony-level benefits may include enhanced disease or parasite resistance and increased productivity due to a varied
workforce with a range of skills.

Q: How was the group-level benefit of genetic diversity within honey bee colonies supported by an experiment?
A: An experiment set up colonies of honey bees with queens inseminated either once or with sperm from ten drones.
Both types of colonies were then infected with a bacterium causing American foulbrood, a larval disease. The study
found that colonies with polyandrous queens exhibited lower disease intensity and had larger and more robust
colonies by the end of summer. The increased genetic diversity within these colonies resulted in fewer infected
workers and slower disease spread.

Q: What mechanism might underlie colony-level immunity in honey bees, according to research led by David Tarpy?
22

A: The research suggests that colony-level disease resistance in honey bees may result from individual differences in
innate immunity, specifically the nonspecific defense mechanisms of the immune system. By analyzing mRNA from
pools of larvae and measuring the expression of genes encoding antimicrobial peptides, the study found that within-
colony variance among samples in the upregulation of hymenoptaecin (but not of abaecin) decreased with increasing
genetic diversity. This indicates that more genetically diverse colonies exhibited less variation in individual immune
responses, suggesting that individual larval immunocompetence alone may explain colony-level immunity in honey
bees and other eusocial insects.

Q: What did Heather Mattila and Thomas Seeley aim to test in their experiment?
A: They aimed to investigate whether the advantages observed in colonies with polyandrous queens were due to the
ability of a genetically diverse worker force to carry out tasks more efficiently.

Q: How did Mattila and Seeley categorize the colonies in their experiment?
A: They categorized the colonies into two groups: those headed by single-insemination queens and those with queens
inseminated by multiple males.

Q: How did Mattila and Seeley ensure that disease did not affect their experiment's results?
A: They kept the colonies free from disease through antibacterial and antiparasitic treatments.

Q: What were some of the parameters monitored in the experiment?

A: Parameters monitored included the total weight of the bees, the foraging rates of workers, and the mean area of
comb in the hive where offspring are reared and food is stored.

Q: What was the outcome of the experiment regarding the survival of colonies?
A: None of the genetically uniform colonies survived through the winter, while about one-fourth of the colonies led by
polyandrous queens survived.

Q: What conclusion did the findings of the experiment suggest?

A: The findings suggested that the genetic diversity of colonies, boosted by queen polyandry, enhances the fitness of
the queen independently of disease resistance.

FIGURE 10.20 Polyandry provides genetic benefits in honey bees. Genetically diverse colonies (the queens had mated
with several males—shown as blue circles) produced on average more comb than did workers in genetically uniform
23

colonies (the queens had mated with only one drone—shown as red squares). (After H. R. Mattila and T. Seeley. 2007.
Science 317: 362–364.)

Q: How are the genetic compatibility hypothesis and the genetic diversity hypothesis related to the idea of inbreeding
avoidance?
A: Both the genetic compatibility hypothesis and the genetic diversity hypothesis may be related to the idea that
females choose extra-pair mates to avoid inbreeding, which could lead to the production of less heterozygous
offspring.

Q: What evidence supports the inbreeding avoidance hypothesis in fairy-wrens?

A: Studies on red-backed fairy-wrens and purple-crowned fairy-wrens support the inbreeding avoidance hypothesis. In
red-backed fairy-wrens, females paired to genetically similar males were more likely to produce offspring sired by
extra-pair males, and those offspring were less inbred (more heterozygous) than within-pair offspring. In purple-
crowned fairy-wrens, incestuous pairs motivated females to engage in frequent extra-pair copulations, resulting in a
higher percentage of extra-pair offspring compared to non-incestuous pairs.

Q: What are some examples of studies that have investigated the indirect genetic benefits of multiple mating by
females?
A: Studies comparing fitness-related traits in extra-pair versus social mates, extra-pair paternity rates in pairs varying
in genetic similarity or inbreeding, and genetic diversity or fitness-related traits in extra-pair versus within-pair
offspring provide examples.

Q: What did an analysis of 16 avian species find regarding the benefits of polyandry for offspring performance traits?
A: An analysis of 16 avian species found that polyandry was not significantly beneficial for any single offspring
performance trait, including growth rate, survival, or adult size.

Q: What did Erol Akçay and Joan Roughgarden find in their analysis of 55 avian species regarding the good genes or
genetic compatibility hypotheses?
A: Erol Akçay and Joan Roughgarden found little support for the good genes or genetic compatibility hypotheses in
more than half of the 55 avian species studied. They observed that extra-pair males were on average larger and older
than within-pair males, but they found no differences in terms of secondary sexual traits, body condition, or genetic
relatedness to the female. Additionally, they found no significant correlation between pair genetic similarity and extra-
pair paternity.

Q: What did another meta-analysis of 39 studies from 33 species of birds find regarding the occurrence of extra-pair
paternity and the relatedness of social mates?
A: Another meta-analysis of 39 studies from 33 species of birds found a positive relationship between the occurrence
of extra-pair paternity and the relatedness of social mates. They also suggested that the positive association may
depend on the type of molecular marker used in the studies, implying that methodological issues could affect the
results of previous studies.

Q: What is the focus of the discussion in the provided text?

A: The focus is on the potential direct benefits of polyandry for females, rather than just indirect genetic benefits.
24

Q: What are some examples of direct benefits of polyandry mentioned in the text?
A: Examples of direct benefits of polyandry include access to additional resources or parental care, better protection
from sexually harassing males, and reduced risk of infanticide via paternity confusion.

Q: What are the hypotheses presented in Hypotheses 10.3 regarding direct benefits of polyandry?
A: The hypotheses presented in Hypotheses 10.3 regarding direct benefits of polyandry are the additional resources
hypothesis and the additional care hypothesis.

Q: Can you explain the additional resources hypothesis and the additional care hypothesis?
A: - Additional resources hypothesis: Females mate polyandrously to gain access to additional resources from their
partner.

Additional care hypothesis: Females mate polyandrously to gain more caregivers to help rear young.

Q: What is the additional resources hypothesis regarding polyandry?

A: The additional resources hypothesis suggests that females mate multiply to gain access to resources necessary for
successful reproduction.

Q: How do female red-winged blackbirds benefit from engaging in extra-pair copulations according to the text?
A: Female red-winged blackbirds may be allowed to forage for food on the territories of males with which they have
engaged in extra-pair copulations, whereas genetically monogamous females are chased away.

Q: How do male butterflies in polyandrous species potentially bribe females into mating with them?
A: Male butterflies in polyandrous species potentially bribe females into mating with them by providing them with
nutrients in the form of spermatophores, which are more nutritious in highly polyandrous species compared to
monogamous species.

Q: How was the prediction regarding reproductive output and polyandry tested in butterfly species?
A: The prediction regarding reproductive output and polyandry was tested in butterfly species by using the mean
number of copulations for females of each species as a measure of its degree of polyandry. Researchers then
quantified reproductive output by measuring the mass of eggs produced by a female in the laboratory and dividing
that figure by the body mass of the female to control for differences among species in weight.
25

FIGURE 10.21 Polyandry boosts reproductive output in pierid butterflies. In eight pierid species, the mean reproductive
output of females (measured as the cumulative mass of eggs produced divided by the mass of the female) increases
with the mean number of mates a female has during her lifetime. (After C. Wiklund et al. 2001. Proc R Soc London B
368: 1661–1667.)

Q: What unique reproductive structure do female Neotrogla barklice possess?

A: Female Neotrogla barklice possess an elaborate, penislike structure called a gynosome.

Q: What is the function of the gynosome in female Neotrogla barklice?

A: The gynosome in female Neotrogla barklice is used to receive a spermatophore from the male and anchor their
bodies together for up to 3 days of copulation.

Q: How do males and females of Neotrogla barklice differ in terms of sexual organs?
A: Females of Neotrogla barklice have a gynosome, whereas males lack an intromittent organ altogether.

Q: Why might multiple mating by females be advantageous in Neotrogla barklice?

A: Multiple mating by females in Neotrogla barklice might be advantageous because it allows them to receive
potentially lifesaving direct benefits in the form of nutritious spermatophores, which are crucial in an environment
where food sources such as bat carcasses and guano are scarce.

Q: How does polyandry in dunnocks enable females to access additional parental assistance?
A: Polyandry in dunnocks enables females to access additional parental assistance by distributing their copulations
between multiple males, ensuring that both sexual partners contribute to the rearing of the brood.

Q: What behavior do female dunnocks exhibit to ensure both males contribute to parental care?
26

A: Female dunnocks actively solicit matings from whichever male, alpha or beta, usually the latter, has had less time
in their company, ensuring that both males reach a paternal threshold necessary for contributing to parental care.

Q: What evidence supports the benefits of polyandry in dunnocks?

A: In a population of introduced dunnocks in New Zealand, polyandrous groups had higher hatching and fledging
success than monogamous pairs, providing evidence for the benefits of polyandry in this species.

FIGURE 10.23 Adjustment of copulation frequency by polyandrous female dunnocks. (A) A female dunnock (B) living in
a territory with two males solicits copulations more often from the male that has spent less time with her, whether that
is the alpha or the beta male. (After N. B. Davies et al. 1996. Anim Behav 51: 27–47.)

Likewise, a female superb starling that is pair-bonded with one male sometimes mates with another unpaired male in
her social group; the second male then sometimes helps rear the brood of his mate (Rubenstein 2007). By copulating
with an additional male, the polyandrous female has in effect made it advantageous for the “other male” to invest
parentally in her offspring. Recall from earlier that female superb starlings also gain indirect genetic benefits in the
form of increased offspring heterozygosity when they mate with males from outside their social group—males that
would be unable to contribute any parental care (Rubenstein 2007). Dunnocks too receive some indirect genetic
benefits in the form of an amelioration of the negative effects of inbreeding (Santos et al. 2015), in addition to the
direct benefits. Together, these studies remind us that females can, and in some species do, receive both direct and
indirect benefits from mating with several males.

Self-Study Questions 10.2

[Please note: You must be using an online, browser-based eReader in order to view this content.]

1. In the wattled jacana, females fight for and defend territories that accommodate several males. A territorial female
then mates with all of the members of her harem, and each male cares for his own clutch of eggs. However, 75
percent of these clutches are of mixed paternity. This example illustrates that in polyandrous systems

males tend to exert more control than females over mating, while females exert more control than males over
parental care.
27

males must engage in extra-pair copulations to gain any reproductive fitness.

to have any reproductive success, males may bear a cost of caring for offspring that are not his own.

sexual selection pressure is generally weak on both males and females.

males that join a female's harem have lower fitness than those that do not.

2. By mating with multiple males, females may gain indirect benefits such as

access to additional resources necessary for successful reproduction.

additional parental assistance in caring for offspring and protecting offspring from infanticidal rivals.

increasing the odds that her partner's genes are identical to her own.

increased control over mating decisions.

increased genetic diversity, resulting in more heterozygous offspring.

3. Which example provides support for the additional resources hypothesis for the evolution of polyandry?

The offspring of female dark-eyed juncos produced with an extra-pair mate have higher reproductive success than
those produced with her pair-bonded mate.

Red-backed fairy-wren females socially paired to genetically similar males are more likely to produce young sired by
extra-pair males.

Female dunnocks copulate with a second, subordinate male when the alpha male is away, and both sexual partners
help her rear her brood.

Female red-winged blackbirds are allowed to forage on territories of males with which they have had extra-pair
copulations.

Genetically diverse bee colonies are able to produce on average more comb (to rear offspring and store food) than
genetically uniform colonies.

10.3 Polygyny: Multiple Mating by Males

LEARNING OBJECTIVES
■ Evaluate how the resource needs and grouping decisions of females dictate male reproductive behavior, leading to
the formation of either female defense or resource defense polygyny.
28

■ Describe the alternative explanations for why males congregate at leks and why females prefer specific lekking
individuals.
■ Review the costs, benefits, and ecological factors that produce scramble competition polygyny.

Q: What are the four hypotheses listed in HYPOTHESES 10.4 to explain why males are polygynous?
A: The four hypotheses are the female defense polygyny hypothesis, the resource defense polygyny hypothesis, the
lek polygyny hypothesis, and the scramble competition polygyny hypothesis.

Q: How does the female defense polygyny hypothesis explain polygyny in males?
A: The female defense polygyny hypothesis suggests that when resources are evenly distributed but females form
groups, males can follow and guard a group of females, gaining access to reproductively ready females more easily.

Q: According to the resource defense polygyny hypothesis, what strategy do males employ to gain access to females?
A: The resource defense polygyny hypothesis posits that when resources are clumped, attract multiple females, and
are easily defended, males guard the resources and the females by setting up territories.

Q: What factors determine the type of polygynous mating system in a species?

A: The type of polygynous mating system in a species is determined by the degree to which females are clustered in
defensible groups as a result of predation pressure or food distribution.

Q: What is the female defense polygyny hypothesis?

A: The female defense polygyny hypothesis suggests that when resources are evenly distributed in space but females
form groups, males will follow and guard a group of females.

Q: According to the resource defense polygyny hypothesis, what do males guard?

A: According to the resource defense polygyny hypothesis, males guard the resources, and by extension, the females,
by setting up a territory.

Q: What does the lek polygyny hypothesis propose?

A: The lek polygyny hypothesis proposes that when resources are distributed heterogeneously and females are
widespread and do not form groups, males will wait for females to come to them.

Q: What does the scramble competition polygyny hypothesis suggest males will do?
A: The scramble competition polygyny hypothesis suggests that when resources are distributed heterogeneously and
females are widespread and do not form groups, males will seek out females.

Female Defense Polygyny

When receptive females occur in defensible clusters, males will compete directly for those clusters and female defense
polygyny will result. For example, nests of Cardiocondyla ants produce large numbers of virgin queens, leading to
lethal combat among the males of the colony, the survivors of which get to copulate with dozens of freshly emerged
females (Heinze et al. 1998). But one does not have to be hyperaggressive to practice female defense polygyny. For
example, males of tiny siphonoecetine amphipods construct elaborate cases composed of pebbles and fragments of
mollusk shells found in the shallow ocean bays where they live. They move about in these houses and capture females
29

by gluing the females’ houses to their own, eventually creating an apartment complex containing up to three potential
mates (FIGURE 10.24) (Just 1988).

Q: What is female defense polygyny, and where is it commonly observed in vertebrates?

A: Female defense polygyny is a mating system where males guard and defend groups of females, particularly in
species where females form groups. It is commonly observed in vertebrates, particularly in mammals.

Q: Why does female defense polygyny occur in species where females form groups?
A: Female defense polygyny occurs in species where females form groups because it makes defending multiple
females at the same time much easier for males.

Q: Can you provide examples of species where female defense polygyny is observed?
A: Yes, examples include bighorn sheep (Ovis canadensis), western gorillas (Gorilla gorilla), and male lions (Panthera
leo). In these species, males compete to control sexual access to groups or harems of females that form for protection
against predators.

FIGURE 10.25 Female defense polygyny in the greater spear-nosed bat (Phyllostomus hastatus). A large male (bottom
right) guards a roosting cluster of smaller females. A successful male may sire as many as 50 offspring with his harem
females in a year.

Q: How do dominant males of the Montezuma oropendola employ female defense polygyny?
A: Dominant males of the Montezuma oropendola employ female defense polygyny by controlling clusters of nesting
females. They shift from nest tree to nest tree, following females rather than defending a nesting resource per se.
30

Q: What is the significance of the behavior observed in Montezuma oropendolas?

A: The behavior observed in Montezuma oropendolas demonstrates that dominant males employ the female defense
tactic rather than a resource defense tactic when the level of competition makes this possible.

Q: Can you describe the mating strategy of dominant males in Montezuma oropendolas?
A: Dominant males secure up to 80 percent of all matings by driving subordinate males away from clusters of nesting
females. They do not defend specific nesting resources but rather follow females to control mating opportunities.

FIGURE 10.26 Female defense polygyny in the Montezuma oropendola. (A) Male Montezuma oropendolas attempt to
monopolize females in (B) small colonies of nesting females. (C) As colony size increases, mating attempts are often
disrupted by rivals. (D) As a result of these disruptions, frequency of copulations per hour at the colony site decreases.
(C, D after M. S. Webster and S. K. Robinson. 1999. Am Nat 154: 717–729.)

Q: What dictates the behavior of males in most polygynous species?

A: In most polygynous species, it is the behavior of females that dictates the behavior of males, ultimately determining
the type of mating system that forms.

Q: How do male plains zebras attempt to defend a harem of females?

A: Male plains zebras attempt to defend a harem of females by forming groups to reduce the risk of predation and to
avoid harassment by bachelor males. This type of female defense polygyny is referred to as harem defense polygyny.

Q: What benefits do female zebras derive from associating with dominant males?
31

A: Female zebras that associate with dominant males are able to feed for 10 percent longer per day compared to
those that associate with subordinate males, as dominant males successfully keep other males away, reducing
harassment and allowing females to forage safely and continuously.

Q: Why do female zebras form groups with dominant males?

A: Female zebras form groups with dominant males because it allows them to forage longer without being harassed by
other males. Additionally, grouping together provides more eyes to spot predators, enhancing their safety while
feeding.
Q: What is resource defense polygyny?
A: Resource defense polygyny occurs when a male becomes polygynous by controlling access to resources that
females visit on occasion, rather than by guarding groups of females.

Q: How does resource distribution influence the mating system in Grevy’s zebra compared to plains zebra?
A: In Grevy’s zebra, where grass and water sources are scarce and more heterogeneously distributed in arid regions,
dominant males set up territories near water where receptive females spend most of their time. This allows them to
mate with the most females by defending the best territories where more females are likely to be. In contrast, plains
zebra males can monopolize both receptive and nonreceptive females by guarding groups for extended periods of
time in regions where resources are more accessible and evenly spread.

Q: What factors influence the differences in group formation and mating systems between plains zebras and Grevy’s
zebras?
A: The differences in group formation and mating systems between plains zebras and Grevy’s zebras are influenced by
the availability and distribution of resources. Plains zebras occur in slightly wetter regions with more accessible and
evenly spread grass and water sources, facilitating the formation of female groups overseen by a male. In contrast,
Grevy’s zebras live in more arid regions with scarce and heterogeneously distributed resources, leading to shorter-
lived female associations and males setting up territories near water sources to mate with receptive females.

Q: What is resource defense polygyny, and how does it manifest in Lamprologus callipterus?
A: Resource defense polygyny occurs when males control access to resources that females need for reproduction. In
Lamprologus callipterus, territorial males defend suitable nest sites and collect empty snail shells from the lake bottom
to create middens containing up to 86 shells. By monopolizing these shell-rich territories, males can attract and mate
with multiple females, as up to 14 females may nest simultaneously in different shells within one male's midden.

Q: How does sexual size dimorphism contribute to the mating system of Lamprologus callipterus?
A: Lamprologus callipterus exhibits extreme sexual size dimorphism, with males being more than 12 times heavier
than females. This size difference allows territorial males to defend nest sites and collect snail shells, which are
essential resources for female reproduction. Female body size is constrained by the size of snail shells, while male
body size is not, enabling larger males to monopolize territories and mate with multiple females.

FIGURE 10.27 Resource defense polygyny in the African cichlid fish Lamprologus callipterus. (A) A territorial male
bringing a shell to his midden. The more shells that there are in a male’s midden, the more nest sites are available for
females to use. (B) The tail of one of the territorial male’s very small mates can be seen in a close-up of the shell
(lower left) that serves as her nest.
32

Q: How does resource defense polygyny manifest in black-winged damselflies (Calopteryx maculata)?
A: Male black-winged damselflies defend floating vegetation that serves as a safe location for females to lay their
eggs. Each female mates with the defending male before depositing her eggs in the vegetation he controls. By
monopolizing this resource, territorial males attract multiple mates.

Q: Can you provide an example of resource defense polygyny in insects other than damselflies?
A: Antlered flies (Phytalmia spp.) engage in resource defense polygyny by fighting for small rot spots on fallen logs or
branches of certain tropical trees. These locations attract sexually receptive females, which mate with successful
territory defenders before laying their eggs. The more of this resource a male holds or the longer he holds it, the more
likely he is to acquire multiple mates.

FIGURE 10.28 Resource defense polygyny in an Australian antlered fly. Males compete for possession of egg-laying
sites that can be found only on certain species of recently fallen trees.

Q: How did Nick Davies and Arne Lundberg alter the mating system of the dunnock?
A: Davies and Lundberg altered the mating system of the dunnock by providing supplemental oats and mealworms to
some females for months at a time. This caused the females' home ranges to contract substantially. As a result,
females with reduced ranges had fewer social mates, leading to a shift from female polyandry to female monogamy.

Q: What did the study by Davies and Lundberg reveal about the relationship between resource distribution and mating
systems?
A: The study showed that altering the distribution of key resources, such as supplemental food for females, can
change the spatial distribution of potential partners. As a female's home range contracted due to the availability of
resources in a smaller area, the capacity of one male to monitor her activities increased. This led to a shift in the
33

mating system from female polyandry to female monogamy, supporting the theory that males attempt to monopolize
females within the constraints imposed by the spatial distribution of resources and potential partners.

FIGURE 10.29 A test of the female distribution theory of mating systems. When food supplementation reduces the size
of a female dunnock’s home range, a male can monopolize access to that female and reduce the number of males
with which she interacts. (After N. B. Davies and A. Lundberg. 1984. J Anim Ecol 53: 895–912.)

Q: What did Stanislav Pribil and William Searcy investigate regarding female red-winged blackbirds?
A: Pribil and Searcy investigated the concept of a "polygyny threshold" in female red-winged blackbirds. They tested
whether females would choose to mate with already mated males if the quality of the territories controlled by those
males was experimentally boosted, while the value of territories controlled by unmated males was lowered.

Q: What did Pribil and Searcy find regarding female mate choice in red-winged blackbirds?
A: Pribil and Searcy found that female red-winged blackbirds, which typically choose unmated males, would reverse
their usual preference if the quality of territories controlled by mated males was enhanced. When provided with the
opportunity to nest over water, even as part of a polygynous male's harem, females reared almost twice as many
young on average compared to females nesting onshore in what became a monogamous male's territory.

Q: Why might some female red-winged blackbirds and females of other avian species accept a second-rate territory,
even though it typically results in fewer offspring?

A: Some female birds, like female red-winged blackbirds, may accept inferior territories because the costs of finding
other potential partners are high, or because the remaining unmated males have extremely poor territories. For some
females, the options may be limited to accepting secondary status or not breeding at all. Unmated female pied
flycatchers, for example, may choose already mated males voluntarily and with full knowledge, possibly because of
the high costs associated with finding other potential partners or because the remaining unmated males have poor
territories.
Q: What role does female-female competition play in the evolution of mating systems, particularly in pied flycatchers?
34

A: Female-female competition for access to mates and resources, although less studied than male-male competition, is
crucial to the evolution of mating systems. In populations of pied flycatchers, nearly half of the females exhibit a white
forehead patch, a trait shared with males. Research indicates that females with patches produce more offspring, are
more aggressive to intruders, and are less likely to be attacked by intruders. These findings suggest that female pied
flycatchers are under strong selection to prevent their mates from accepting additional females onto their territories,
underscoring the differing sexual interests between males and females in polygynous mating systems.

FIGURE 10.30 Female pied flycatchers with white forehead patches aggressively deter rivals from their mate’s
territory. (A) An adult pied flycatcher female without forehead patch and (B) an adult pied flycatcher female with a
forehead patch (left) and a male showing the typical breeding plumage, including the forehead patch (right).

Q: What is lek polygyny?

A: Lek polygyny is a mating system where males do not defend groups of females or resources; instead, they establish
small display arenas where females come to choose mates.

Q: What do males do in lek polygyny?

A: In lek polygyny, males fight to control small areas used exclusively as display arenas to attract females.

Q: What is the purpose of leks in lek polygyny?

A: Leks serve as locations where females come to choose mates, receive sperm, and then depart to rear offspring
alone.

Q: How are leks established?

A: Leks may be clustered in traditional display grounds or scattered in dispersed leks. They are often located where
females pass through or where it's convenient for displaying to females.

Q: Can you provide an example of a species that uses lek polygyny?

A: Yes, the white-bearded manakin is an example. In Trinidadian forests, female manakins visit leks to choose mates
from numerous males performing elaborate displays.

Q: What is a standard feature of lekking species in terms of male mating success?

A: Huge inequalities in male mating success, known as high reproductive skew, are standard features of lekking
species.

Q: How do older males in topi antelopes benefit in a lek?

A: In topi antelopes, older males tend to occupy the center of a lek, resulting in more copulations compared to younger
rivals forced to the periphery.

Q: How do females in topi antelopes behave in regards to mating with preferred males?
35

A: Females in topi antelopes prefer males in the center of the lek and are willing to compete aggressively with other
females for matings with these preferred males on central lek territories.

Q: What percentage of males in a lek of West African hammer-headed bats are responsible for the majority of
matings?
A: Only 6 percent of the males in a lek of West African hammer-headed bats are responsible for 80 percent of the
matings.

Q: Describe the mating behavior of West African hammer-headed bats.

A: In West African hammer-headed bats, males gather in groups along riverbanks, defend display territories high in
trees, and produce loud cries resembling "a glass being rapped hard on a porcelain sink" to attract receptive females,
who visit several males before mating.

FIGURE 10.31 Mating success at topi leks. Each male topi in a central position at his lek mates with more females per
capita than do males forced to peripheral sites. The dark green bars represent the averages for the three leks shown.
Bars depict mean ± SE. (After J. Bro-Jørgensen and S. M. Durant. 2003. Anim Behav 65: 585–594.)

FIGURE 10.32 A lek polygynous mammal: the hammer-headed bat. Males cluster in dense groups at mating arenas
where they defend display territories in trees overhanging riverbanks.

Q: Why do male hammer-headed bats exhibit lekking behavior?

36

A: Bradbury argues that lekking behavior in male hammer-headed bats evolves when other mating tactics do not pay
off due to the wide and even distribution of females. Hammer-headed bats travel over great distances in search of
widely scattered food sources, making it difficult for males to monopolize females directly or indirectly. Thus, males
display their merits to choosy females that come to leks to inspect them.

Q: How is the absence of defensible clusters of females or resources related to lek polygyny?
A: The absence of defensible clusters of females or resources has been proposed as an explanation for both
nonterritorial scramble competition polygyny and lek polygyny. Species facing these conditions may either search for
mates individually or form elaborate leks, but why some species choose one strategy over the other is not entirely
clear.

Q: What are some potential reasons why males of lekking species congregate in traditional leks rather than displaying
solitarily in dispersed leks?
A: One possibility is that congregated males in traditional leks may be relatives that assist one another in mate
attraction. However, evidence for male relatedness driving lekking behavior is limited. Another possibility is that
predation favors the adoption of a safety-in-numbers tactic among congregated males, but this idea has been rarely
tested.
Q: What are three hypotheses related to female choice of top males in lekking species?
A: According to the hotspot hypothesis, males cluster in places ("hotspots") where the routes frequently traveled by
receptive females intersect. The hotshot hypothesis argues that subordinate males cluster around highly attractive
males to have a chance to interact with females drawn to these "hotshots." Lastly, the female preference hypothesis
suggests that males cluster because females prefer sites with large groups of males, where they can more quickly or
safely compare the quality of many potential mates.

Q: How does the hotspot hypothesis explain the clustering of males in lekking species?
A: The hotspot hypothesis suggests that males congregate in places where the routes frequently traveled by receptive
females intersect, creating areas of high female traffic. Males cluster in these "hotspots" to increase their chances of
encountering receptive females.

Q: What is the hotshot hypothesis regarding male clustering in lekking species?

A: The hotshot hypothesis proposes that subordinate males cluster around highly attractive males, known as
"hotshots," to have a chance to interact with females drawn to these top males. By associating with hotshots,
subordinate males may increase their own chances of mating.

Q: How does the female preference hypothesis explain male clustering in lekking species?
A: The female preference hypothesis suggests that males cluster because females prefer sites with large groups of
males. This allows females to more quickly or safely compare the quality of many potential mates, leading to
increased efficiency in mate selection.

Q: What is the hotspot hypothesis in the context of male congregation at leks?

A: The hotspot hypothesis suggests that males congregate in places, known as "hotspots," where the routes frequently
traveled by receptive females intersect.

Q: According to the hotshot hypothesis, how do subordinate males behave in relation to highly attractive males?
37

A: The hotshot hypothesis proposes that subordinate males cluster around highly attractive males, known as
"hotshots," to have a chance to interact with females drawn to these top males.

Q: What does the female preference hypothesis suggest regarding male clustering at leks?
A: The female preference hypothesis suggests that males cluster because females prefer sites with large groups of
males. This allows females to quickly or safely compare the quality of many potential mates.

Q: How did Frédéric Jiguet and Vincent Bretagnolle test the alternative hypotheses for why males congregate at leks?
A: They created artificial leks populated by plastic decoys resembling males and females of the little bustard bird
species and observed the response of living little bustards to these decoys.

Q: What did the experiment by Jiguet and Bretagnolle reveal about the hotspot hypothesis?
A: The experiment showed that female decoys failed to attract males, leading to the rejection of the hotspot
hypothesis.

Q: How did the experiment support the hotshot hypothesis?

A: Male decoys, especially those resembling individuals with highly symmetrical plumage patterns, regularly attracted
both females and males, supporting the hotshot hypothesis.

Q: What additional finding from the experiment suggested support for the female preference hypothesis?
A: The experiment found that clusters of four decoys attracted more females per decoy than smaller groups, which is
consistent with the female preference hypothesis. However, adding more than four decoys did not draw in additional
females.

Q: How did a study of the great snipe contribute to testing the hotspot and hotshot hypotheses?
A: In the study of the great snipe, removal of central dominant males caused neighboring subordinates to leave their
territories, supporting the hotshot hypothesis. However, removal of a subordinate while the alpha snipe was in place
resulted in quick replacement on the vacant territory by another subordinate, indicating that the presence of attractive
hotshots, not the real estate per se, determines where clusters of males form.

Q: What hypothesis has been supported by evidence in some cases regarding male lekking behavior?
A: The hotspot hypothesis has received support in certain cases regarding male lekking behavior.

Q: Can you provide an example of evidence supporting the hotspot hypothesis?

A: Yes, for example, a site where male fallow deer gathered to display shifted when logging activity altered the paths
regularly followed by fallow deer of both sexes, indicating support for the hotspot hypothesis.

Q: How do Indian peacocks support the hotspot hypothesis?

A: Indian peacocks tend to gather near areas where potential mates are feeding, known as hotspots. The removal of
some males has no effect on the number of females visiting leks of this species, suggesting that females are
inspecting those areas because of the food they provide, not because of the males there.

Q: What evidence supports the hotspot hypothesis in the case of certain tropical manakins?
38

A: The leks of certain tropical manakins are located in areas relatively rich in fruits, the primary food source of these
birds. This is consistent with the idea that males gather near places with resources attractive to females, supporting
the hotspot hypothesis.

FIGURE 10.33 A test of the hotspot hypothesis. (A) The location of greater sage-grouse (Centrocercus urophasianus)
leks (numbered black circles) in relation to sagebrush, meadows, forests, and a lake. (B) The distribution of nesting
females in relation to the leks where males gather to display. The darker the shading, the more females present.
Launch the Interactive Figure (After R. M. Gibson. 1996. Anim Behav 52: 993–1005.)

Q: Why does the hotspot hypothesis not apply to ungulates like the Kafue lechwe?
A: The hotspot hypothesis does not apply to ungulates like the Kafue lechwe because ovulating females leave their
customary foraging ranges to visit groups of males some distance away, possibly to compare the performance of
many males simultaneously.

Q: What is the female preference hypothesis regarding the Uganda kob?

A: According to the female preference hypothesis, a female preference for quick and easy comparisons might make it
advantageous for males of the Uganda kob to form large groups.

Q: What was observed regarding the operational sex ratio across leks of different sizes in the Uganda kob?
39

A: Contrary to the prediction based on the female preference hypothesis, the operational sex ratio is the same for leks
across a spectrum of sizes in the Uganda kob, indicating that males are no better off in large groups than in small
ones.

Q: How does the barking treefrog challenge the female preference hypothesis?
A: In the case of the barking treefrog, the more males chorusing in a pond, the more receptive females show up on a
given night. However, preventing males from calling at the pond does not reduce the number of females arriving,
which goes against the expectation that a large number of calling males is needed to attract a large number of choosy
females.

FIGURE 10.34 Female density in the Kafue lechwe is not correlated with lek formation. Four leks of the antelope (open
circles) are not located in the areas of highest female density, providing evidence against the hotspot hypothesis for
this species. Launch the Interactive Figure (After A. Balmford et al. 1993. Behav Ecol Sociobiol 33: 57–65.)

FIGURE 10.35 Female Uganda kob do not aggregate disproportionately at leks with large numbers of males. (A)
Female attendance at leks is simply proportional to the number of males displaying there. (B) As a result, the female-
to-male ratio does not increase as lek size increases. (After J. C. Deutsch. 1994. Behav Ecol Sociobiol 34: 451–459.)
40

FIGURE 10.36 The relationship between mating females and calling males of the barking treefrog. The more female
barking treefrogs mating at a pond on a given night, the more males found chorusing there. However, the relationship
stems not from the ability of large numbers of males to attract more females with their calls but from the fact that
both sexes respond similarly to a set of environmental variables, including temperature and rainfall. (After C. G.
Murphy. 2003. Behav Ecol 14: 274–281.)

Q: Why does no single hypothesis for why lekking males form groups apply to every species?
A: No single hypothesis for why lekking males form groups applies to every species because the reasons for lek
formation can vary across different species.

Q: What do interactions among males on a lek usually enable?

A: Interactions among males on a lek usually enable individuals of high physiological competence to demonstrate their
superior condition to their fellow males and to visiting females.

Q: Why might it be advantageous for females to come to a lek?

A: It might be advantageous for females to come to a lek in order to compare and choose a partner of the highest
quality, as lekking males are forced to compete in ways that separate the strong from the weak.

Q: Has the possibility of showy females in some species following similar rules been explored?
A: Yes, the possibility of showy females in some species following similar rules has been explored in at least one sex-
role reversed species.

Q: What tactic do males employ in species where receptive females and resources are widely dispersed?
A: In species where receptive females and resources are widely dispersed, males employ a tactic called scramble
competition polygyny.

Q: How do flightless females of Photinus fireflies affect male behavior?

A: Flightless females of Photinus fireflies affect male behavior by appearing almost anywhere over wide swaths of
Florida woodland, prompting males of this species to search widely for them rather than establishing territories.

Q: What determines mating success in Photinus fireflies?

A: In Photinus fireflies, mating success is determined by the persistence, durability, and perceptiveness of males in
finding females rather than by aggression.

Q: How do male thirteen-lined ground squirrels search for females?

A: Male thirteen-lined ground squirrels search widely for females, which become receptive for only 4 to 5 hours during
the breeding season, and the first male to find an estrous female and copulate with her will fertilize about 75 percent
of her ova.

Q: What special kind of intelligence might male fitness depend on in thirteen-lined ground squirrels?
A: Male fitness in thirteen-lined ground squirrels might depend on the ability to remember where potential mates can
be located, as returning males often revisit places where they have interacted with about-to-become-receptive
females.
Q: How did Andrew Sih and colleagues test the prediction regarding the mating system in water striders?
41

A: Andrew Sih and colleagues experimentally altered pool and group size in water striders and followed the same
males in both treatments to observe changes in mating behavior.

Q: What mating system did males exhibit in small groups in large pools, and how did it differ from their behavior in
small pools with large groups?
A: In small groups in large pools, males exhibited scramble competition, whereas in small pools with large groups, they
often exhibited female defense polygyny, where the largest male drove other males into hiding but allowed females to
be relatively active.

Q: How did matings differ between the two experimental conditions in water striders?
A: In large pools with small groups, copulations typically lasted for nearly two hours, while in small pools with large
groups, they were less frequent and much shorter in duration.

Q: Why did the researchers hypothesize that matings were shorter in small pools with large groups of water striders?
A: The researchers hypothesized that matings were shorter in small pools with large groups of water striders because
the dominant male had less need to mate repeatedly and little need to guard females after transferring sperm due to
driving off competitors.

Q: How does the breeding behavior of male wood frogs differ from that of other species with explosive breeding
assemblages?
A: Male wood frogs eschew territorial behavior and instead hurry about trying to encounter as many egg-laden females
as possible before the one-night orgy ends, as females are available only on that one night.

FIGURE 10.37 An explosive breeding assemblage. A male wood frog grasps a female (upper left) that he has found
before rival males, two of which are near the mating pair. Numerous fertilized egg masses float in the water around
the frogs.

Self-Study Questions 10.3

1. The main factor that differentiates female defense polygyny from resource defense polygyny is
42

the distribution of females and resources; in female defense polygyny, resources are evenly distributed but females
form defensible groups. correct

sexual selection pressure; in female defense polygyny, there is stronger selection pressure on males, leading to male-
male competition. incorrect

parental care; in female defense polygyny, high predation risk necessitates male parental care. incorrect

infanticide risk; in female defense polygyny, the grouping of females leads to a higher risk of infanticide by rival males.
incorrect

female choice; in female defense polygyny, females are choosy, whereas in resource defense polygyny males are
choosy. incorrect
the distribution of females and resources; in female defense polygyny, resources are evenly distributed but females
form defensible groups.

Section reference: 10.3 Polygyny: Multiple Mating by Males

Topic: LO1 Evaluate how the resource needs and grouping decisions of females dictate male reproductive behavior,
leading to the formation of either female defense or resource defense polygyny.

2. In an experiment in the great snipe, removing the central dominant male from the lek caused neighboring
subordinate males to leave the territory. However, when a subordinate male was removed, he was quickly replaced by
another subordinate. This example illustrates

the hotspot hypothesis, where males cluster in places where females frequently travel. incorrect

the hotshot hypothesis, where males cluster around highly attractive males to have a chance to interact with females.
correct

the female preference hypothesis, where males cluster because females prefer sites with large groups of males.
incorrect

a polygyny threshold, where females gain more benefits by mating with a polygynous male than by not. incorrect

harem defense polygyny, where females are socially bonded in harems that are easily monopolized by males.
incorrect

Section reference: 10.3 Polygyny: Multiple Mating by Males

Topic: LO2 Describe the alternative explanations for why males congregate at leks and why females prefer specific
lekking individuals.

3. Scramble competition polygyny is likely to occur when

43

there is little sexual selection pressure on males, so male-male competition and reproductive skew are low. incorrect

resources are clumped but females are widespread and don't form groups. incorrect

there is a high cost to defending a territory, and females are unlikely to form groups, search for males, or frequent the
same place. correct

there is high predation pressure, causing females to form groups to dilute predation risk. incorrect

there is an absence of defensible clusters of females or resources, and females search for males with whom to mate.
incorrect

Section reference: 10.3 Polygyny: Multiple Mating by Males

Topic: LO3 Review the costs, benefits, and ecological factors that produce scramble competition polygyny.there is an
absence of defensible clusters of females or resources, and females search for males with whom to mate.

10.4 Polygynandry and Promiscuity: Multiple Mating by Both Sexes

LEARNING OBJECTIVES
■ Identify the different associations that distinguish polygynandrous and promiscuous mating systems.
■ Review examples of polygyandry, considering the ecological and social conditions that underlie this form of mating
system.
■ Describe the roles of parental care and sexual conflict in the evolution of promiscuity.

Q: What is the term used by behavioral biologists to describe a mating system where both sexes have multiple
partners?
A: Behavioral biologists refer to this type of mating system as polygynandry or promiscuity.

Q: What is the distinction between polygynandry and promiscuity?

A: Polygynandry refers to cases where there are pair-bonds between males and females that tend to live together in
multi-sex groups, whereas promiscuity describes cases when mating occurs in the absence of long-term relationships.

Q: Which type of mating system, polygynandry or promiscuity, is considered to be more common?

A: Promiscuity is considered to be more common, particularly in species that do not provide parental care and those
with external fertilization, such as many fishes.

Q: What is polygynandry?
A: Polygynandry is a mating system where males and females form pair-bonds and tend to live together in multi-sex
groups.

Q: In what types of species does polygynandry tend to occur?

A: Polygynandry tends to occur in species that form social groups.

Q: Can you provide an example of a species exhibiting polygynandry?

44

A: One example of a species exhibiting polygynandry is the dunnock.

Q: How does the mating system vary within dunnock populations?

A: Within dunnock populations, the mating system can vary, including monogamy, polyandry, polygyny, and
polygynandry.

Q: What factors contribute to the variation in mating systems within dunnock populations?
A: The variation in dunnock mating systems is thought to be a result of resource availability and sexual conflict.

Q: What species besides the dunnock exhibits polygynandrous groups?

A: The alpine accentor (Prunella collaris) is another species that forms polygynandrous groups.

Q: How do alpine accentors typically form polygynandrous groups?

A: Alpine accentors typically form polygynandrous groups with one dominant male, one to three subordinate males,
and two to four females.

Q: What environmental factors contribute to the formation of polygynandrous groups in alpine accentors?
A: The alpine accentors live at high elevations and feed on widely dispersed and patchily distributed insects, leading to
overlapping ranges in both males and females, which contributes to the formation of polygynandrous groups.

Q: What is the role of parental care in polygynandrous groups of alpine accentors?

A: Both dominant and subordinate males actively engage in parental care at nests in polygynandrous groups of alpine
accentors.

Q: How does resource availability influence grouping patterns in species like the dunnock and the alpine accentor?
A: The density of both birds and resources drives grouping patterns in species like the dunnock and the alpine
accentor.
Q: How does the mating behavior of the great tinamou differ from that of dunnocks and alpine accentors?
A: The great tinamou exhibits promiscuity, where females mate with multiple males who then provide care for the
combined clutches of eggs. Unlike dunnocks and alpine accentors, multiparent care is unnecessary in the great
tinamou because its young are precocial and can largely take care of themselves after hatching.

Q: Describe the reproductive strategy of the great tinamou.

A: In the great tinamou, females roam the forest floor in search of males and may mate with up to three males. After
mating, females leave the males with the clutch of eggs, and it is the fathers that incubate the combined clutches of
eggs.

Q: What distinguishes the promiscuity of Littorina saxatilis from that of dunnocks and alpine accentors?
A: In Littorina saxatilis, both males and females mate indiscriminately and frequently, with males siring clutches of
offspring without any involvement in parental care. This differs from dunnocks and alpine accentors, where
multiparent care is necessary for provisioning food to young.

Q: Explain the parental care behavior in Littorina saxatilis.

45

A: In Littorina saxatilis, females provide care by retaining fertilized eggs inside a brood pouch until they hatch. Only
the mother takes care of the young, while males do not contribute to offspring care despite mating with multiple
females.
Q: Describe the mating behavior of the seaweed fly Coelopa frigida.
A: In Coelopa frigida, males and females exhibit promiscuous mating behavior without any courtship rituals. Males
attempt to mount females whenever they encounter them, leading to frequent mating attempts. Females often
acquiesce to these attempts, resulting in high mating frequencies.

Q: What is convenience polyandry, and how does it relate to the mating behavior of Coelopa frigida?
A: Convenience polyandry refers to a mating strategy where females mate with multiple males without active mate
choice or courtship behavior. In Coelopa frigida, females often acquiesce to male mating attempts, leading to repeated
matings with multiple males, which may be a consequence of sexual conflict between males and females.

Q: How does male harassment influence the mating behavior of Coelopa frigida?
A: Male harassment is common in Coelopa frigida, with males frequently attempting to mount females. This results in
promiscuous mating behavior, where females mate multiple times with different males, potentially as a strategy to
minimize the negative effects of male harassment or to ensure fertilization by multiple males.

FIGURE 10.38 Seaweed flies exhibit extreme promiscuity. When population densities are high, both males and females
may copulate hundreds of times during their short, 3-week lives.

Self-Study Questions 10.4

1. How does polygynandry differ from promiscuity?

Polygynandry is driven by the need for parental care, while promiscuity is driven by the distribution of individuals and
resources. incorrect

Polygynandry requires pair-bonds and occurs in species with social groups, while promiscuity occurs without long-term
relationships. correct

Polygynandry occurs when females or resources are widely distributed, while promiscuity occurs when females and
resources are clustered. incorrect

Polygynandry evolves as a result of female choice, while promiscuity evolves when neither sex is able to monopolize
the other. incorrect

They are interchangeable terms for mating systems in which both sexes mate multiply. incorrect
46

Section reference: 10.4 Polygynandry and Promiscuity: Multiple Mating by Both Sexes
Topic: LO1 Identify the different associations that distinguish polygynandrous and promiscuous mating systems.

2. Alpine accentors form polygynandrous groups with one dominant male, up to three subordinate males, and two to
four females. They feed on insects that are patchily distributed over the landscape, leading to overlapping ranges in
males and females. Reproduction and parental care are shared by males and females. This example illustrates that

polygynandry evolves as a result of the constraints imposed by choosy females on males. incorrect

the distribution of resources is the key determinant in the formation of groups that leads to polygynandrous mating
systems. incorrect

polygynandry is maladaptive for both males and females and is the result of neither sex being able to gain an
advantage over the other. incorrect

the density of both individuals and resources, as well as the need for parental care, facilitates grouping and multiple
mating in both sexes. correct

mating systems such as polygynandry are determined by ecological and social circumstances and are thus unlikely to
change once established. incorrect

Section reference: 10.4 Polygynandry and Promiscuity: Multiple Mating by Both Sexes
Topic: LO2 Review examples of polygynandry, considering the ecological and social conditions that underlie this form
of mating system.

3. Promiscuity is most common in species exhibiting

high reproductive skew for males. incorrect

long-term social relationships or pair-bonds. incorrect

dense clustering of both males and females. incorrect

extreme sexual dimorphism between males and females. incorrect

little to no parental care by either parent. correct

Section reference: 10.4 Polygynandry and Promiscuity: Multiple Mating by Both Sexes
Topic: LO3 Describe the roles of parental care and sexual conflict in the evolution of promiscuity.
47

SUMMARY
1.Mating systems can be defined in terms of the number of sexual partners an individual acquires during a breeding
season. Monogamy occurs when males and females both have only a single mate. Polygyny occurs when males have
multiple partners but females have just one, whereas polyandry occurs when females have multiple partners and
males have just one. When both sexes mate multiply, it is termed polygynandry if there is a pair-bond between males
and females, and promiscuity if there is no association between sexual partners beyond sperm transfer.

2.Monogamy is likely to evolve when the costs to mating multiply are prohibitively high for both sexes and neither sex
can monopolize access to mates or resources. This can occur because potential mates do not form groups and roam
widely, making them potentially costly to locate, when individuals have the ability to restrict mating behavior in their
partner, or when resources are so critical to successful reproduction that both parents are necessary to rear young.

3.Polyandry is likely to evolve when the benefits of mating multiply are higher for females than remaining
monogamous. These potential benefits include indirect benefits in the form of genetic advantages for offspring or
direct benefits for the female. Potential genetic benefits include higher individual offspring quality or viability, greater
genetic compatibility between the female and her mate, and greater genetic diversity among a female’s offspring.
Females also may mate with many partners to avoid the risk of inbreeding. Potential direct benefits include access to
additional resources, access to additional parental care, better protection from sexually harassing males, and reduced
risk of infanticide.

4.The diversity of polygynous mating systems has evolved in response to different patterns of female distribution,
which are often driven by the distribution of resources or the risk of predation. These female decisions in turn affect
the profitability of different kinds of mating tactics that males use to gain access to females. When females or the
resources that they need are clumped in space, female defense or resource defense polygyny becomes more likely. If,
however, females are widely dispersed or male density is high, males may engage in nonterritorial scramble
competition for mates, or they may acquire mates by displaying at a lek where females come specifically to seek out
mates and nothing else.
5.Multiple mating by both sexes is likely to evolve when neither sex is able to gain an advantage over the other.
Species living in social groups, often because of resource or offspring-care needs, are likely to form long-term
relationships and to be polygynandrous when multiple caregivers are required to successfully raise young. In contrast,
when there is no long-term association between males and females and often uniparental or no form of parental care,
both sexes may mate promiscuously, or even indiscriminately. Promiscuous mating systems are often the result of
strong sexual conflict between males and females.

Chapter 10 Flashcards

Quiz

Test Bank
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Multiple Choice
1. Sexual selection differs from natural selection in that
a. the traits under sexual selection can be environmentally influenced, unlike those that are the
product of natural selection.
b. hereditary differences among phenotypes are required for natural selection to occur, but not
for sexual selection to occur.
c. sexual selection can produce traits that harm the survival of the species, whereas natural
selection acts to preserve the species over the long term.
d. natural selection affects a broader spectrum of traits than sexual selection, which acts only on
individual differences in the ability to gain access to mates.
Answer: d

Questions 2−3.
Females of a species of fly form swarms in which several dozen individuals gather together to fly
about in circles. When a male enters the swarm, females inflate their abdomens and move toward
the male, who carefully selects one of the many females as his partner.

2. What is there about this species that constitutes a Darwinian puzzle?

a. The means by which females manage to enlarge their abdomen, which requires a special and
still-mysterious abdomen-inflating device
b. The fact that the members of one sex have evolved a truly bizarre display to induce the
opposite sex to mate with them
c. The fact that females have to persuade apparently choosy males to mate with them
d. The fact that many females have to wait for a mate, which reduces the reproductive potential
of the species
Answer: c

3. Which is an example of a group selectionist puzzle?

a. The means by which females manage to enlarge their abdomen, which requires a special and
still-mysterious abdomen-inflating device
b. The fact that the members of one sex have evolved a truly bizarre display to induce the
opposite sex to mate with them
c. The fact that females have to persuade apparently choosy males to mate with them
d. The fact that many females have to wait for a mate, which reduces the reproductive potential
of the species
Answer: d

Questions 5−9.
Kirk’s dik-dik is a small African antelope that usually forms male–female pairs.

4. Is it possible that monogamy evolved in this species because males are attempting to guard
their partners from other males? This is an example of
a. a hypothesis.
b. a prediction.
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c. test evidence.
d. a scientific conclusion.
Answer: a

5. Males place their scent marks over those of the female, which hides the signals of sexual
receptivity that females produce when they are fertile. This is an example of
a. a hypothesis.
b. a prediction.
c. test evidence.
d. a scientific conclusion.
Answer: c

6. It may be that if males were experimentally removed from pairs, females would wander from
their territories into the territories of other males. This is an example of
a. a hypothesis.
b. a prediction.
c. test evidence.
d. a scientific conclusion.
Answer: b

7. If DNA data were collected from the offspring of intact pairs, it should be found that the
offspring of the females were sired by their social partners.
Hypothesis This is an example of
a. a hypothesis.
b. a prediction.
c. test evidence.
d. a scientific conclusion.
Answer: b

8. It is also possible that pairs form in order to protect their offspring, with two adults doing a
better job of protection than just one. This is an example of
a. a hypothesis.
b. a prediction.
c. test evidence.
d. a scientific conclusion.
Answer: a

9. Genetic monogamy is
a. extremely rare.
b. the rule (with a few exceptions) among mammals, but not among birds.
c. the rule (with a few exceptions) among birds, but not among mammals.
d. never found in reptiles.
Answer: a
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10. Male ostriches guard a nest into which a female lays an egg from their mating. The next day
she mates with a different male; the first male also mates with a different female, which places
her egg in his nest. This species exhibits
a. polygynandry.
b. polyandry.
c. monogamy.
d. polygyny.
Answer: a

11. If you study a pair of European beavers, you will find that all of the young in their colony are
genetic offspring of the pair. This species exhibits
a. polygynandry.
b. polyandry.
c. monogamy.
d. polygyny.
Answer: c

12. Long-term storage of sperm in some species of tortoise allows a female to produce a clutch
of eggs with offspring fathered by multiple males. This species exhibits
a. polygynandry.
b. polyandry.
c. monogamy.
d. polygyny.
Answer: b

13. Red deer males compete to control access to an entire herd of does, meaning that a single
male will mate with multiple females, while other males have zero fitness. This species exhibits
a. polygynandry.
b. polyandry.
c. monogamy.
d. polygyny.
Answer: d

Short Answer
14. Give the primary reason that monogamy in male animals would be considered a Darwinian
puzzle.
Answer: Males generally have sufficient sperm to fertilize a great many eggs, which ought to
favor males that mate with a great many egg producers.

15. Polyandry in females might be advantageous if polyandrous females were to secure some
genes from their partners that were especially compatible with the genes in their eggs. State a
prediction that follows from this hypothesis.
Answer: (Answers may vary.) In order to expand the potential gene pool, the mating partners of
females will vary substantially from female to female.
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16. Why have behavioral ecologists concluded that the ecology of females has more to say about
the evolution of the mating system of a species than the ecology of males?
Answer: Receptive females are usually in short supply, not the other way around.