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CHAPTER 7: The Evolution of Cooperation

MULTIPLE CHOICE

1. By definition, altruistic behaviors

a. incur a cost to the recipient.
b. incur a cost to the actor.
c. result in a benefit to the actor.
d. are beneficial to the recipient and the actor.
ANS: B DIF: Easy REF: Altruism: A Puzzle
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Applying

2. Altruism at first was a puzzle to evolutionary biologists because

a. it should be more common in nature because it increases the fitness of recipients.
b. it should be more common in nature because it increases the fitness of species.
c. it should not be common in nature because it decreases the fitness of actors.
d. it should not be common in nature because it decreases the fitness of species.
ANS: C DIF: Medium REF: Altruism: A Puzzle
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding

3. Which of the following is true of altruism?

a. Altruism is a strategy favored by sexual selection.
b. Altruism can evolve only if the benefit to the group is very high compared with the cost to
the actor.
c. Altruism cannot simultaneously increase the fitness of all members of the group.
d. Altruism cannot evolve unless there is no cost to the actor.
ANS: B DIF: Medium REF: Altruism: A Puzzle
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding

4. Two unrelated male baboons work together to guard a female in estrus to keep away a more dominant
male. This is an example of
a. mutualism. c. reciprocal altruism.
b. altruism. d. kin selection.
ANS: A DIF: Easy REF: Mutualism
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Applying

5. Mutualistic behaviors
a. are common in nature when animals work together.
b. are rare because “slacking” is often profitable for individuals, not groups.
c. involve only kin and never unrelated individuals.
d. are profitable for the actor but not the recipient.
ANS: B DIF: Medium REF: Mutualism
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding
 6. Coalitions among male baboons
a. allow middle-ranking, cooperative males a chance to gain access to receptive females.
b. are an example of mutualistic behavior as all males get equal mating opportunities.
c. do not allow lower-ranking males to outcompete higher-ranking males.
d. can predict male grooming relationships.
ANS: A DIF: Hard REF: Mutualism
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding

7. Group selection
a. is likely to occur in primates because they live in well-established social groups.
b. is likely to occur in primates because certain behaviors benefit the whole group.
c. is unlikely to occur in primates because certain behaviors may benefit the group to the
detriment of the actor.
d. is unlikely to occur in primates because it is a theoretically unsound principle.
ANS: C DIF: Hard REF: The Problem with Group-Level Explanations
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding

8. Although alarm calling benefits the whole group, it cannot be explained by group selection because
a. callers make themselves conspicuous to the predators, but calling costs little in terms of
individual fitness.
b. calling reduces the risk of mortality for everyone who hears the call, changing the
frequency of callers and noncallers in the population, because everyone benefits.
c. noncallers benefit from the alarm call and will have higher fitness than the callers, so
selection will suppress alarm calling.
d. callers and noncallers have the same relative fitness.
ANS: C DIF: Hard REF: The Problem with Group-Level Explanations
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Understanding

9. When group selection and individual selection are opposed,

a. group selection prevails if groups are large and there is little migration.
b. individual selection prevails.
c. group selection usually prevails because the conditions for individual selection are too
stringent.
d. group selection never prevails.
ANS: B DIF: Hard REF: The Problem with Group-Level Explanations
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding

10. The key to the evolution of altruism is that

a. recipients have to be more likely to carry the altruistic allele than nonrecipients.
b. actors have to be more likely to carry the altruistic allele than nonactors.
c. recipients have to be unrelated to the actors.
d. nonrecipients have to be more likely to carry the altruistic allele than recipients.
ANS: A DIF: Easy REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Remembering
11. Altruistic behavior is more likely
a. among kin than among nonkin.
b. among nonkin than among kin.
c. among kin than among nonkin, but only when kin and nonkin have the opportunity to
interact.
d. among nonkin than among kin, but only when nonkin interact with each other more than
kin do.
ANS: A DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Remembering

12. Kin selection predicts

a. that group selection is more likely in families.
b. that altruism is more likely among relatives.
c. that altruism should occur only in higher animals.
d. that only 50% of related individuals are altruistic.
ANS: B DIF: Easy REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Applying

13. Hamilton’s rule states that

a. altruism evolves only among nonkin.
b. selfish genes swamp altruistic genes.
c. altruistic behavior is favored if the cost to the actor is less than the benefit to the recipient,
devalued by the degree of relatedness.
d. altruistic behavior is favored if the benefits to the actor are greater than the costs to the
recipients, devalued by their degree of relatedness.
ANS: C DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Remembering

14. The coefficient of relatedness (r) is a measure of

a. each individual’s fitness benefits.
b. the probability that two individuals acquire the same allele through descent from a
common ancestor.
c. the probability that two randomly chosen individuals share an allele.
d. sibling relatedness.
ANS: D DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Remembering

15. The coefficient of relatedness (r) between full siblings is

a. 0.25. c. 0.75.
b. 0.5. d. 1.00.
ANS: B DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Applying

16. The coefficient of relatedness (r) between mother and son is

a. 0.25. c. 0.75.
 b. 0.5. d. 1.00.
ANS: B DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Applying

17. Imagine that an alarm caller sacrifices its life to save other conspecifics. According to Hamilton’s rule,
how many full siblings would it have to save for the behavior to be favored?
a. at least 1. c. at least 3.
b. at least 2. d. Hamilton’s rule cannot be satisfied.
ANS: C DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Applying

18. Imagine a caregiver who helps raise a conspecific’s offspring. Such caregiving reduces her fitness by
50% and increases the fitness of the conspecific by 50%. According to Hamilton’s rule, in which of the
following scenarios could this behavior evolve?
a. Groups of full siblings c. Both types of groups
b. Groups of half siblings d. Neither type of group
ANS: D DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Applying

19. Imagine a caregiver who helps raise a conspecific’s offspring. Such caregiving reduces her fitness by
10% and increases the fitness of the recipient by 25%. According to Hamilton’s rule, in which of the
following scenarios could this behavior evolve?
a. Groups of full siblings c. Both types of groups
b. Groups of half siblings d. Neither type of group
ANS: A DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Applying

20. If the coefficient of relatedness between two individuals is zero (r = 0), then
a. altruism can evolve if c > b.
b. altruism can evolve if c < b.
c. altruism cannot evolve via kin selection.
d. altruism will sometimes evolve regardless of the values of c and b.
ANS: C DIF: Medium REF: Kin Selection
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Applying

21. If the coefficient of relatedness between two individuals is 0.5, then

a. altruism can evolve if c > 2b.
b. altruism can evolve if c < 2b.
c. altruism cannot evolve via kin selection.
d. altruism will sometimes evolve regardless of the values of c and b.
ANS: B DIF: Medium REF: Hamilton’s Rule
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Applying

22. Which of the following is predicted by Hamilton’s rule?

 a. No altruism should evolve when r < 0.5.
b. Altruism evolves only when r > 0.0.
c. Altruism evolves only when r < 0.0.
d. No altruism should evolve when r > 0.5.
ANS: B DIF: Medium REF: Hamilton’s Rule
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Understanding

23. Primate altruism includes

a. grooming conspecifics. c. autogrooming.
b. sibling rivalry. d. sharing home ranges.
ANS: A DIF: Easy REF: Kin Biases in Behavior
OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Remembering

24. Phenotypic matching

a. is context dependent.
b. is common among apes but not monkeys.
c. is the ability to recognize kin by a feature such as their smell.
d. favors maternal kin over paternal kin.
ANS: C DIF: Easy REF: Kin Recognition
OBJ: Discuss the mechanisms that allow primates to recognize their relatives.
MSC: Remembering

25. Recent studies on kin recognition have found that age may provide a good proxy measure of paternal
kinship in
a. monogamous species.
b. species where one male dominates mating activity in a group.
c. groups with polygynous mating strategies.
d. polyandrous groups.
ANS: B DIF: Medium REF: Kin Recognition
OBJ: Discuss the mechanisms that allow primates to recognize their relatives.
MSC: Remembering

26. Monkeys and apes recognize their kin through

a. genotype mapping.
b. patterns of male and female associations.
c. contact with their mothers.
d. age estimation of adult males in the group.
ANS: C DIF: Easy REF: Kin Recognition
OBJ: Discuss the mechanisms that allow primates to recognize their relatives.
MSC: Remembering

27. Studies of kin recognition by female rhesus monkeys suggest that

a. females had weak affinities for their maternal half sisters.
b. facial resemblance could be a cue females use to distinguish paternal half sisters.
c. females preferred agemates to maternal and paternal half sisters.
d. females preferred maternal half sisters to unrelated agemates.
ANS: B DIF: Hard REF: Kin Recognition
OBJ: Discuss the mechanisms that allow primates to recognize their relatives.
 MSC: Remembering

28. When monkeys recognize paternal kin, they may rely on

a. gender. c. age similarity.
b. genotype. d. phenotype.
ANS: C DIF: Medium REF: Kin Recognition
OBJ: Discuss the mechanisms that allow primates to recognize their relatives.
MSC: Remembering

29. Baboon females

a. have strong affinities for both maternal and paternal half sisters.
b. are not able to distinguish maternal and paternal half sisters from unrelated females.
c. prefer maternal half sisters over paternal half sisters.
d. prefer paternal half sisters over maternal half sisters.
ANS: A DIF: Medium REF: Kin Recognition
OBJ: Discuss the mechanisms that allow primates to recognize their relatives.
MSC: Remembering

30. Kin selection combined with females remaining in their natal group, in many primate species, has led
to
a. mutualistic behavior.
b. asymmetrical behavior.
c. the transmission of rank laterally from sister to sister.
d. the transmission of rank from mothers to offspring.
ANS: D DIF: Medium REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Remembering

31. The apparent stability in female dominance relationships in female-bonded primates may be due to
a. abundant food supply. c. sexual selection.
b. kin selection. d. habitat seasonality.
ANS: B DIF: Medium REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Understanding

32. Which of the following is usually more common among kin than among nonkin in primate groups?
a. Aggression c. Alarm calling
b. Foraging d. Coalition formation
ANS: D DIF: Easy REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Remembering

33. Which of the following describes maternal rank and matrilineages in macaque, vervet, and baboon
groups?
a. Maternal rank is transferred to offspring, particularly daughters.
b. Maternal kin occupy dissimilar ranks in the dominance hierarchy.
c. Ranking within matrilineages is usually laterally transferred.
d. Female dominance relationships are unstable over time.
ANS: A DIF: Medium REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
 MSC: Remembering

34. Which of the following statements do studies of macaques, vervets, and baboons support?
a. Maternal rank is not a good predictor of a daughter’s rank.
b. Matrilineages rank above or below all members of other matrilineages.
c. The stability of female dominance relationships is not a result of kin-based alliances.
d. Females inherit patrilineal rank.
ANS: B DIF: Medium REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Remembering

35. Studies of red howler monkeys have shown that coalitions of

a. fathers and daughters form enduring bonds that are the basis for social groups.
b. related males are more likely to experience rank reversals.
c. unrelated males last four times as long as coalitions of unrelated males.
d. fathers and their maturing sons are necessary to repel incursions into their group by alien
males.
ANS: D DIF: Medium REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Remembering

36. The cooperative breeding system of primates such as marmosets and tamarins can be explained by
a. chimerism, which increases fraternal twins’ inclusive fitness when they help raise one
another’s offspring.
b. helpers who care for the offspring of the breeding pair, although the helpers are usually
not related to them.
c. the fact that mothers sometimes allow their daughters to breed.
d. mutualism between the nonbreeding helpers.
ANS: A DIF: Hard REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Understanding

37. Parent–offspring conflict occurs

a. because parents and offspring share all of their genes.
b. because individuals are more closely related to self than to siblings.
c. only when siblings are not full siblings.
d. only during weaning.
ANS: D DIF: Medium REF: Parent–Offspring Conflict
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Understanding

38. Which of the following is true regarding parent–offspring conflict?

a. Weaning is an example of parent–offspring conflict, and this transition reflects a
fundamental symmetry in the genetic interests of mothers and offspring.
b. A mother is equally related to each of her offspring; in contrast, an offspring is at most
25% related to its siblings but is 100% related to itself.
c. A mother is motivated to wean her current infant so she can begin investment in her next
infant, but the infant will resist its mother’s attempts at weaning because it is not in its
genetic best interest.
d. Parent–offspring conflict only occurs in primates because reproductive costs are higher
than for other mammals.
 ANS: C DIF: Medium REF: Parent–Offspring Conflict
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Understanding

39. Which of the following is predicted to occur after conflict results in violence as a way to mend
relationships?
a. Huddling c. Coalition formation
b. Reconciliation d. Altruism
ANS: B DIF: Easy REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Applying

40. Monkey A lives in a group with full siblings. Monkey B lives in a group with half siblings. Monkey C
lives in a group with cousins. And monkey D lives in a group with its grandparents. For which monkey
in which group is altruistic behavior most likely to occur?
a. A c. C
b. B d. D
ANS: A DIF: Medium REF: Hamilton’s Rule
OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Applying

41. Of the following, which is the most certain way to identify kin in primates?
a. Phenotypic matching c. Proximity/context
b. Age/cohort d. Reconciliation
ANS: C DIF: Easy REF: Kin Recognition
OBJ: Discuss the mechanisms that allow primates to recognize their relatives.
MSC: Understanding

42. Matrilines are an example of

a. altruism. c. grooming relationships.
b. kin recognition. d. dominance relationships.
ANS: B DIF: Easy REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Applying

43. Even primate “friends” sometimes come into conflict. Their aggressive interactions may not have a
detrimental effect on social cohesion if there is
a. kin selection. c. grooming.
b. altruism. d. reconciliation.
ANS: D DIF: Easy REF: Kin Biases in Behavior
OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Applying

44. In most promiscuously mating primate species that live in multimale, multifemale groups, mothers
share 50% of their genes with their offspring. Different offspring of the same female are likely to share
what percentage of their genes with each other?
a. 0% c. 50%
b. 25% d. 100%
ANS: B DIF: Medium REF: Parent–Offspring Conflict
 OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Applying

45. Reciprocal altruism requires

a. many interactions between kin.
b. the ability to count.
c. that there are no slackers or cheaters.
d. sufficient memory to keep track of altruistic and nonaltruistic acts.
ANS: D DIF: Easy REF: Reciprocal Altruism
OBJ: Evaluate arguments about the importance of reciprocal altruism in primate groups.
MSC: Understanding

46. Evidence for reciprocal altruism

a. is absent among primates because they cannot recognize kin.
b. is absent among primates because they do not have long-term memories.
c. exists among primates because they often live in stable social groups.
d. exists among primates because of their longevity.
ANS: C DIF: Easy REF: Reciprocal Altruism
OBJ: Evaluate arguments about the importance of reciprocal altruism in primate groups.
MSC: Understanding

47. In a study done with vervets, researchers found that

a. genetic relatedness is necessary for contingent reciprocity to occur.
b. they responded to recruitment calls from other monkeys after receiving grooming bouts
from them.
c. contingent reciprocity can evolve as confederates react appropriately to the presence and
absence of reciprocity.
d. agemates were more likely to be reciprocal.
ANS: C DIF: Hard REF: Reciprocal Altruism
OBJ: Evaluate arguments about the importance of reciprocal altruism in primate groups.
MSC: Remembering

48. In recent studies of chimpanzees, researchers found that

a. possessors of food were less likely to share their food with individuals who had recently
groomed them.
b. grooming is not reciprocal.
c. grooming is sometimes a precursor for food sharing.
d. grooming is exchanged for mating opportunities.
ANS: C DIF: Medium REF: Reciprocal Altruism
OBJ: Evaluate arguments about the importance of reciprocal altruism in primate groups.
MSC: Remembering

49. Reconciliatory contact between opponents immediately following an aggressive encounter can reduce
rates of self-scratching, an indicator of stress. If no reconciliation is attempted, then we can assume
that
a. rates of self-scratching drop back to baseline, regardless of stress level.
b. rates of self-scratching remain above baseline, indicating high stress levels.
c. only one opponent experiences high stress levels.
d. the opponent will have high rates of aggression between them in the future.
ANS: B DIF: Medium REF: Reciprocal Altruism
 OBJ: Evaluate arguments about the importance of reciprocal altruism in primate groups.
MSC: Applying

ESSAY

1. Compare altruism and mutualism in terms of the fitness effects on actors and recipients.

ANS:
Altruistic acts provide fitness benefits to the recipient but at a fitness cost to the actor. Mutualistic acts
provide fitness benefits to both the recipient and the actor but only under strict conditions that prevent
slackers or cheaters from benefiting at the expense of the other party. Cooperative action by both
individuals must be required for success.

DIF: Easy REF: Altruism: A Puzzle | Mutualism

OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding

2. What is altruism? Why was it a puzzle for evolutionary biology before Hamilton?

ANS:
Altruism is behavior that benefits another at a cost to the actor. As students have learned so far, natural
selection will favor only traits that benefit the individual actor, and therefore, since these acts are a
detriment to the actor’s reproductive success, they never should evolve and should in fact be selected
against. Hamilton demonstrated how these behaviors could be selected for if they are directed toward
kin because they share some percentage of each other’s genes. It would have been nice if the authors
had actually introduced the term “inclusive fitness” here, and instructors may wish to do so.

DIF: Medium REF: Altruism: A Puzzle

OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding

3. What is group selection? Why do evolutionary biologists believe that individual selection will be more
powerful than group selection when the two are in conflict?

ANS:
Group selection is the idea that behaviors that benefit the group as a whole will be selected even if
they result in declines in fitness for the actor. This concept is problematic for evolutionary biologists
for several reasons. First, a behavior that benefits the group but to the detriment of the actor could
theoretically disappear, as those that have risky behaviors have reduced reproductive success and
therefore have fewer offspring who will perform that behavior in the next generation. Thus, selection
at the level of the individual should be stronger. Furthermore, for group selection to work, there would
need to be extensive variation between groups, as there is for individuals, but that does not appear to
be the case.

DIF: Medium REF: The Problem with Group-Level Explanations

OBJ: Understand why altruism is unlikely to evolve in most circumstances.
MSC: Understanding

4. What is Hamilton’s rule? What are the two fundamental predictions of Hamilton’s rule? Devise a
primate scenario in which these two predictions of Hamilton’s rule would be satisfied.

ANS:
 Hamilton’s rule is the mathematical model that helps predict whether altruistic behaviors will be
selected for. The equation rb > c specifies that an altruistic behavior will be selected for when r, the
coefficient of relatedness (the likely proportion of genes shared between two individuals), multiplied
by b, the fitness benefits for the recipients of such behavior, is greater than c, the fitness cost to the
actor. The two predictions that follow from Hamilton’s rule are that (a) altruism should be directed to
kin only because r = 0 for nonkin and (b) closer kinship allows for more costly altruism because r is
higher and can offset greater costs to the actor. There is extensive evidence within primates that known
or supposed kin, but especially known maternal kin, perform more altruistic behaviors such as
grooming, supporting each other in coalitions, food sharing, cooperative defense of territories and
groups, and helping for cooperatively breeding species.

DIF: Hard REF: Kin Selection

OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Analyzing

5. Construct a hypothetical example to illustrate how a gene causing altruism in an individual could
increase in frequency through kin selection.

ANS:
The example used repeatedly in the book is the clearest illustration of how a gene causing altruism in
an individual could increase in frequency through kin selection. Imagine a monkey species that has a
special alarm call for birds of prey. This alarm call is genetically determined, so individuals who carry
the “raptor alarm call gene” warn their group that there is predation threat. Since they share genes with
their kin, including the “raptor alarm call gene,” if they reside in a group with kin they help them
survive when they call, even though they make themselves more obvious to the bird of prey. As
generations go by, the genes shared by relatives keep being passed on and in this way may spread.

DIF: Medium REF: Kin Selection

OBJ: Understand how evolution can favor altruism through the processes of kin selection and
reciprocal altruism. MSC: Analyzing

6. What is the evidence for maternal and paternal kin recognition? Illustrate your answer with examples.

ANS:
Maternal kin selection is straightforward: Individuals can learn by proximity and familiarity that the
offspring of their mother are their kin. In primates, maternity is the one certainty. Therefore, it is not
surprising to find evidence of nepotism in female primates that reside in their natal groups with their
mothers, such as baboons, macaques, and vervet monkeys: Female kin inherit their rank from their
mother, have adjacent rank to their sisters, form matrilines that allow these large-scale coalitions to
compete with other such matrilines within their group, and groom each other more frequently.
Paternity is harder to determine in primates in promiscuously mating species. There do seem to be
some trends, however. Dominant males tend to sire more of the offspring in multimale, multifemale
groups, and so paternity can be assumed. For these species, age may be a proxy for paternity. But even
so, some species, such as macaques, seem to have preferences for paternal kin. Female macaques have
more affiliative interactions with paternal half siblings so they may be able not only to distinguish age
but also to use some other cue, perhaps a phenotypic one. Male primates may also be able to recognize
siblings, as the example of red howler males cooperatively defending groups demonstrates.

DIF: Medium REF: Kin Selection

OBJ: Discuss the mechanisms that allow primates to recognize their relatives.
MSC: Applying

7. Conflict and aggression are a large part of group life for many primate species, which would seem
disruptive to social bonds. What, then, keeps social groups cohesive?
 ANS:
Social cohesion in primate groups is achieved through affiliative interactions. Grooming is the number
one social currency in primates. It establishes social bonds and keeps individuals in proximity to one
another. It also functions to decrease the physiological effects of stress. But just because there are
affiliative behaviors does not mean that aggression does not occur. It has been documented that
reconciliation after aggression helps several haplorrhine species maintain positive affiliative
interactions and maintain social cohesion even after aggressive interactions. Furthermore, at least one
sex often resides in a group with kin, and kin selection may also influence social bonds to some
degree. Although the text does not specifically point this out, a student would not be incorrect to draw
on this information, as it is discussed for the majority of the chapter.

DIF: Easy REF: Reciprocal Altruism

OBJ: Evaluate arguments about the importance of reciprocal altruism in primate groups.
MSC: Understanding

8. What evidence exists for reciprocal altruism in primates? Why would reciprocal altruism be common
for primates but not for other animals?

ANS:
Reciprocal altruism would be likely for primates but not other animals because it requires advanced
cognitive skills that may be absent in other species. Relative brain size, our proxy for cognitive skills,
is also large in only a few other mammals. Evidence from primates includes grooming and food
sharing in chimpanzees, vervet coalitionary support and grooming, male chimpanzee coalitionary
support, and food sharing.

DIF: Easy REF: Reciprocal Altruism

OBJ: Evaluate arguments about the importance of reciprocal altruism in primate groups.
MSC: Remembering

9. Explain the phenomenon of genetic chimeras and how it relates to kin selection in cooperatively
breeding marmoset groups.

ANS:
Genetic chimerism is a rare phenomenon in nature, but it occurs in marmosets, who frequently give
birth to fraternal twins. Chimerism occurs when each twin carries its own DNA in addition to its
twin’s DNA, thus having two complete sets within its own body. The chimerism extends to all bodily
tissues, including gametes, so that each twin could pass along the genetic material of itself or its twin
to offspring, effectively increasing relatedness among nonbreeding helpers and infants. This could be
the reason we see this kind of cooperation in marmosets.

DIF: Easy REF: Kin Selection

OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Understanding

10. Interpret the following inequality as it pertains to parent–offspring conflict. Apply Hamilton’s rule,
from the current infant’s perspective, assuming a gene expressed by the current infant increases
maternal investment by a small amount.

1.0  (increase in fitness of current fetus)

0.5  (decrease in fitness of future offspring)

ANS:
In the top position is the infant in question, who is related to itself (1.0) and to its sibling in the bottom
position (0.5). Kin selection will increase investment by the mother until the incremental advantage of
another unit of investment in the current infant is twice the cost to future full siblings of the fetus (four
times the cost for half siblings). Asymmetries in genetic expression then lead to a conflict of interest
between mothers and their offspring. Selection favors mothers that provide less investment than their
infants desire, and selection favors offspring that demand more investment than their mothers are
willing to give. This conflict of interest manifests in weaning tantrums and sibling rivalries.

DIF: Hard REF: Kin Selection

OBJ: Explain how kinship influences the distribution of altruism in primate groups.
MSC: Applying
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