Social Behavior in Mammals
The complexity of social behavior might be expected to increase as organisms move from
simple to more sophisticated, but some invertebrates, such as the Portuguese man-of-
war and ants form colonies of individuals that cooperate much more extensively than
mammals do.
A society is a group of individuals of the same species that is organized in a cooperative
manner, typically extending beyond sexual and parental behavior.
The Sociality Spectrum
The same species interact, including mating systems, parent-offspring interactions,
competition, and cooperation among related or unrelated individuals.
In its simplest definition, social means group living, but this includes a wide range of
variation from mammals that live in family units consisting of parents and offspring (e.g.,
North American beavers, to assemblages numbering in the millions (e.g., nursery colonies
of Mexican free-tail bats.
At one end are species that are typically solitary as adults. This includes many carnivores
(e.g., bears, cats, skunks) and omnivores (e.g., opossums, armadillos), as well as
herbivores (e.g. pocket gophers, rhinos, sloths).
However, even mammals that are considered solitary often form temporary associations
with other individuals, for example between mothers and offspring or mates during
breeding.
Figure 23.1 Sociality spectrum. Sociality varies across taxa and sometimes throughout the life of
a mammal. Patterns of sociality range from (A) solitary (e.g., wolverines) to (B) group living, as in
prairie dogs, to (C) highly social (eusocial) as exhibited by the subterranean naked molerat.
At the opposite end of the spectrum are species that always live in groups and that
would be unlikely to survive away from the group. This extreme sociality, seen most
often in insects of the Order Hymenoptera, is referred to as eusocial.
Among mammals, only two species of African mole-rats , the naked mole rat and the
Damaraland mole-rat, exhibit true eusociality.
Why Mammals Live in Groups
Living in social groups is often assumed to be superior to living a more solitary life, yet
costs and benefits are associated with each.
BENEFITS
Protection Against Predators
First, detection of and communication about danger are more rapid when individuals
are in groups.
According to the “many eyes” hypothesis, individuals in large groups need to spend less
time watching for predators and consequently can spend more time on other activities
such as feeding.
Second, if a predator is going to kill one animal, then being in a larger group reduces
each individual’s chances of being the one killed; this is known as dilution of risk.
Individuals could reduce their “domain of danger” by associating with others. called this
the “selfish herd” effect and that an animal’s position in a group could reduce exposure to
predators, such that individuals surrounded by others would be safer, and that the safest
place to be in the center of the group.
being in a group also can reduce an individual’s chances of being killed by a predator by
making it more difficult for the predator to focus on one animal; the result is predator
confusion.
Not all prey animals are passive in response to predators, and some mammals can and
do mount effective counterassaults.
This antipredator strategy is also used in defense of vulnerable young mammals. Adult
musk oxen, which live on the open treeless tundra, form a defensive perimeter around
calves in response to attacks by wolves and other predators.
Finding and Obtaining Food
Sharing information about food resources (whether intentionally or not) is believed to favor
the evolution of sociality. For diurnally foraging animals, this might involve watching
conspecifics in the group but for nocturnal foragers like bats, information about food
resources might be obtained by listening to auditory cues produced by foraging individuals.
studied the lesser bulldog bat, which lives in social groups and forages on flying insects
over water. by eavesdropping on the sounds made by foraging group members, the bats
appeared to forage more efficiently.
cultural transmission of information about seasonal food resources and migratory routes
influenced the evolution of migratory behavior in moose and bighorn sheep. Such
mechanisms are likely important across many terrestrial and marine mammals
that exhibit migratory behaviors.
sociality can also expand foraging options if group-living animals can access food that
might be avoided by solitary individuals. For example, bison are more likely to graze in
open meadows when in larger groups because of the antipredator benefits of grouping.
Similarly, carnivores are sometimes able to hunt larger, more agile, or more dangerous
prey when doing so in a group.
Wolves and African lions are able to take large species of prey—such as bison in the case
of wolves and African buffalo in the case of lions—that would be nearly impossible to kill
alone.
in group hunts, individual female African lions typically form one side of a “winging”
hunting approach, and similar hunting-role specialization has been documented in
bottlenose dolphins in which one individual drives fish toward a group of closely-spaced
“barrier” individuals.
foraging or hunting in groups, animals can reduce the probability that others will steal
their food. Such theft (known as kleptoparasitism) can be carried out by
conspecifics or by other species.
Indeed, kleptoparasitism may sometimes influence sociality in sympatric carnivores
such as cheetahs, African lions, spotted hyenas, and wild dogs.
Opportunities for Social Learning and Culture
Social learning is the process of acquiring knowledge by observation of or interaction with
another individual, usually of the same species.
suggested as important for mammals in general, and primates and cetaceans in particular. it
requires a long period of physiological and psychological dependence. Large-brained and
highly social species, such as dolphins and primates, spend as much as 25% of their lives
dependent on parents or other relatives.
An experimental demonstration of how social learning could affect foraging decisions
comes from a study of wild vervet monkeys in Africa.
The experimenter colored the distasteful corn with different dye… and this consequently
avoided that coloration.
A second example involving a cultural change in feeding behavior involves humpback
whales feeding on small fish in the Gulf of Maine. one whale was observed using a new
technique associated with the typical pattern of bubble feeding.
Other Potential Benefits
for mammals that form social groups include reduction in energy expenditure associated
with thermoregulation via huddling-- for non hibernating small mammals.
free-ranging vervet monkeys demonstrated that during cold winter months, individuals
with more social partners experienced smaller daily fluctuations in body temperature
and higher minimum temperatures (i.e., became less hypothermic) than did animals with
fewer social partners.
Social groups also provide opportunities for gaining access to other important resources
(e.g., nests, burrows, potential mates).
COSTS
Intraspecific Competition for Resources
Competition for food and other resources (e.g., resting sites, mates, shelters) can
increase with group size, resulting in a potential reduction in fitness or increased effort
to attain resources.
primate species found that distance traveled each day increased as a function of group
size and that larger groups spent more time feeding, presumably because of faster
depletion of food resources and competition within the group.
That study also revealed a negative effect on female fecundity, as have other primate
studies, which reported reduced reproductive rates and slower growth of infants in
larger groups.
Spread of Parasites and Diseases
studies support the relationship between parasite infections and host density or group size.
confirmed a positive relationship between sociality and the intensity and prevalence of
contagious parasites across diverse taxa, including mammals.
Other Potential Costs
larger groups can be more obvious to predators, which could counter the benefits of
group size with respect to predation risk. Aggressive encounters among conspecifics are
likely to
be elevated in larger groups, thereby resulting in higher levels of stress, energetic
expenditures, and the potential for injury.
Parental care misdirected to non offspring and killing of young by nonparents are other
potential negative consequences of living in groups.
Brazilian free-tailed bats roost in caves in dense colonies containing millions of bats. Mothers
returning from a night’s foraging for insects have to find their own infant among the
thousands present. Most of the time, they find their own young by vocalization, but 17% of
the time mothers nursed offspring other than their own.
Female white-footed mice with young are aggressive toward strange adults.
in the absence of the mother, intruders usually kill the pups.
How Social Behavior Evolves
A behavior is considered a social behavior when it has fitness consequences for two or
more individuals. Behaviors that increase fitness for both the actor and recipient are
considered mutualistic.
Behaviors that cost the actor but benefit the recipient are classified as altruistic. such
as alarm-calling to warn conspecifics about danger, providing care for unrelated
young, or sharing food.
These acts may appear to be altruistic but could actually confer direct fitness
benefits to the actors. For example, caring for unrelated young allows inexperienced
individuals to gain parenting skills that enhance their ability to raise their own
offspring—a direct fitness benefit to the actor. Helping sisters, brothers, or cousins.
INDIVIDUAL VERSUS GROUP SELECTION
A basic element of sociality is cooperation. Individuals work together, often sacrificing
personal gain, to achieve a common goal that benefits the social group. Subsequent
biologists assumed that groups with cooperating individuals would be more successful
than those without cooperators—a type of group selection.
COOPERATION AMONG KIN
In its simplest form, Hamilton’s kin selection theory suggests that if a gene that causes
altruistic behavior appears in the population, the gene’s success depends ultimately not
on whether it benefits the individual carrying the gene but on the gene’s benefit to
itself. If the individual that benefited by the act is a relative of the altruist and therefore
more likely than a nonrelative to be carrying that same gene, the frequency of that gene
in the gene pool increases.
Figure 23.4 Cooperative behaviors. Many groupliving mammals perform cooperative behaviors such as group hunting,
communal rearing of offspring, and collective defense against predators or conspecific competitors. Such cooperative
behaviors are common among (A) social canids such as wolves (Canis lupus), (B) have been well studied in members in of
the Family Herpestidae (e.g., meerkats), and (C) characterize longlived species like cetaceans (e.g., orcas).
Examples of kin selection have been widely documented in mammals. Sharing prey
among related individuals (e.g., killer whales) or providing warning calls to relatives
when predators are nearby are examples of behaviors that benefit kin at a potential
cost to the actor. A uniquely mammalian example of kin selection is the nursing of non
offspring (allonursing) by related mothers.
Among gray mouse lemurs in Madagascar, adult females typically form small daytime
sleeping clusters. These groups consist of close female relatives that regularly groom and
nurse offspring other than their own and adopt related dependent young after their
mother’s death. among the highly social meerkats of the Kalahari Desert, about half of
the litters born are allonursed, and being a close relative of the biological mother is one of
the factors important in predicting the likelihood of allonursing.
COOPERATION AMONG NON-KIN
cooperation among unrelated individuals. formulated the concept of reciprocal
altruism—“You scratch my back, and I’ll scratch yours.” both the actor and recipient are
simultaneous in mutually beneficial behaviors.
The factors that affect that likelihood are
(1) length of life span—long-lived organisms have a greater chance of meeting again
to reciprocate;
(2) dispersal rate—low dispersal rate increases the chance that repeated interactions
will occur; and
(3) mutual dependence—clumping of individuals, as occurs when avoiding predation,
increases the chances for reciprocation.
A classic example of reciprocity in mammals was first described in common vampire
bats. At night, these bats feed on blood, primarily from cattle and horses, and then
return to a hollow tree to roost during the day. Given their high metabolic rate,
individuals that do not obtain a blood meal within three days will starve, but most
bats that do feed take in more than they need and will sometimes regurgitate blood
into the mouths of roost mates that did not feed that evening.
