Lecture 11

Social Behavior
Territory – exclusive area of a fixed size that is continuously
defended for some period of time through displays or other
acts of defense.

Territorial defense is a flexible and dynamic behavior

Average size of territories is significantly correlated with the

body size of a species

Territory size often differ within species in relationship to prey

availability and social dominance
 Density & Territories
Huxley (1934) proposed that bird territories could be likened to 'rubber
discs.' In other words, as population density increases, territories (like
rubber discs) are compressed (reduced in size) but only to a certain point.
So, there is a limit to the number of territories in an area and, as a result, a
limit to the number of breeding birds in that area. If correct, then there may
be, in any given area, some birds without territories.

Floaters

Several studies with a variety of species have revealed that males (and
females) removed from their territories are usually replaced by
conspecifics (and, often, replaced very quickly). This indicates that there
are 'floaters' (birds without territories) in at least some populations and
suggests, as proposed by Huxley, that territorial behavior may limit
population density.
Fretwell & Lucas
 floaters

Reproductive success

Survival
 Site Dominance
Birds that are currently at a site or bred
at a site in the previous year are almost
always able to regain their territories.

The reason why birds are able to exhibit

site dominance is largely unknown

One potential reason is that they know

the area and are better able to defend it

Another reason may be they have more

at stake and therefore are more willing to
fight for the territory
 Dominance

Older birds are often dominant

Males are often dominant over females

Status signaling is a way in which dominant individuals can coexist with

subordinate without constant physical altercations

Subordinate birds may have to take more chances

 Dominance
There are examples of defenses of other species or mates

Mate guarding is often observed when there is an excess of males

Some species will protect other species in order to gain so

advantages in terms of foraging (e.g. Gulls and Eiders)
 Flocking
Birds flock for protection (more eyes)

Bird flock to help locate food

Most birds form flocks when resources become less
stable and more patchy

Flocks range from loose temporary aggregations to organized

associations of diverse species

Stable flock membership facilitates personal recognition among

individuals and the development of a dominance hierarchy
Horn’s principle of group feeding. If food is more or less evenly distributed
through the environment and can be defended economically it is
energetically most efficient to occupy exclusive territories. But if food occurs
in unpredictable patches, the individuals should collapse their territories to
roosting spots or nest sites and forages as a group
 Why Flock
Competition for resources does not logically promote flocking

Cooperative feeding (e.g. Pelicans)

Beater effect (e.g. African Hornbills)
Poor condition individuals may at least find some food
More eyes to warn other about predators
Predators are less likely to kill a bird in a flock
Safety allows for more time feeding
 Flocking
Harris’s Hawk hunt in teams
which increases there chances of
acquiring prey
 Flocking

Size of the pigeon flock

 Mobbing Predators
Birds often vocally and sometimes physically attack owls,
hawks, and snakes

Drive away enemy

Refine an individual’s ability to identify a predator
Learn the alarm cal of other species
Mix species flocks
Titmice, chickadees, etc
Tropical ant followers

Nuclear species
Sentinels
 Coloniality
Costs of a colony are universal, however the advantages are not

Costs:
increased competition for nest sites
stealing of nest materials
increased physical interference
increased competition for mates
increased competition for food (&, with large colonies, possible depletion of
food resources)
large groups may attract predators & facilitate spread of parasites & diseases
Misdirected parental care (e.g. Cliff Swallows)
Intraspecific brood parasitism (e.g. Cliff Swallows)
 Mating System
Mating systems result for the extent to which one sex
(usually males) can monopolize the other (usually females),
which is determined by the distribution of females, the
amount of parental care needed, phylogenetic constraints,
and mate choice

Monogamy – the prolonged and essentially exclusive pair

bond with a single member of the opposite sex for purposes
of raising young (92% of all bird species)

Polygamy – pair bonds with multiple mates of the opposite

sex (3% of all bird species)

Polygyny – kind of polygamy in which males pairs with two or

more females
 Mating System
Polyandry – a kind of polygamy in which a female pairs with
several males (<1% of all bird species; e.g. phalaropes)

Polygyandry – kind of polygamy in which a females pairs with

several males, each of which also pairs with several different
females. (<1% of all birds species; e.g. tinamous, dunnocks,
ratites)

Promiscuity – indiscriminate sexual relationships usually of

brief duration (6% of all bird species; e.g. grouse,
hummingbirds, manakins)
 Monogamy
Female distribution may promote monogamy

Pair bonds extent form one breeding season to life

Most monogamous birds share responsibility for building the

nest, incubating, and feeding the young

Male participation is essential for successfully raising young

Males cannot monopolize resources necessary for

supporting extra mates
Because of the investment in the partner there is often
substantial evaluation of potential mates
Judge based on ability to raise young
 Monogamous Terns
The ability of a male to provide fish to
the female may signal that the male
will be a good provider for the young

Male feeding of the female may effect

when the eggs are laid and the size of
the eggs
 Extra-pair Copulations
EPCs have been reported in monogamous species, polygynous species, &
polyandrous species

EPCs are copulations with individual(s) other than a mate or social partner.

EPCs may result in extra-pair fertilizations which, in turn, may result in extra-pair
young (EPY).

The percentage of EPY in populations may range from 0 to more than 50%. In
many songbird populations, the percentage of extra-pair young has been found to
be about 10 - 25%, suggesting that at least some individuals in a population benefit
from EPCs:
 EPCs
Benefits of EPCs
Males Females
Increased fitness Fertility insurance
Possible future mate acquisition Genetically diverse young
Insurance against mate's infertility Improved genetic quality
of young
Access to resources
Costs of EPCs
Males Females
Sperm depletion & ejaculate Male retaliation
Increased risk of cuckoldry Risk of injury
Reduction in parental care Harassment from extra-pair males
Increased likelihood of divorce

Cuckoldry is when a male helps raise offspring that are not his
Mate guarding by male Bank Swallows. Males chase other females several
days before their mates begin egg-laying (when their mates are not fertile).
When their mates are fertile (within a few days of laying their first egg),
males chase their own females almost exclusively. Then, once egg-laying
is nearly finished (and mates are no longer fertile), males again chase
other females. Open circles = males chasing a mate; closed circles =
males chasing females other than their mate (From Beecher and Beecher
1979).
 Waterfowl

Socially monogamous
Pair on the wintering grounds
Males do not help raise young
Skewed operational sex ratio
Female natal philopatry
Forced extra-pair copulations
Males may protect females on wintering grounds
 Polygyny
Resource defense polygyny
Males defend areas of habitat where the females want to
nest

No or little male parental care

In North America, 14 of 278 breeding songbird species (11

of which nest in marshes or grasslands) are polygynous
Yellow-headed Blackbirds
Red-winged Blackbirds

Female defense polygyny

Yellow-billed Caciques
Female
fitness

A male's territory quality will be correlated with his mating success

Polygyny should be more common in patchy environments (where

there is more variation in territory quality; 13 of 14 occur in marsh or
grassland habitats
 Polyandry

Females compete for mates

Replacement clutches reproduced rapidly

Spotted Sandpipers

Females are larger and sometimes have secondary sexual

characteristics

Differences in males and females are mediated by hormones

(prolactin testosterone)

Still a lot of unanswered questions about polyandry

 Leks

Males compete for 'status' in communal displays (leks) & females

choose among males

Females get only sperm from males (they raise young elsewhere
unaided by males)

Much variation in male mating success (one or a few males copulate

with many females; other males with none)

Hot spot leks – leks located where females home ranges overlap
(Sage Grouse)

Hop shot – leks form around preferred males (Cock of the Rock)

Conspicuousness – leks form where visible (Ruff)

Cooperative – leks form through cooperation (Long-tailed Manakin)

Why?

Predation
Comparison between males
Learning by males
Cooperation by benefit young males