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Note: In general, terms have been dened as they apply to birds. Nevertheless, many terms (especially those naming basic anatomical structures or biological principles) apply to a range of living things beyond birds. In most cases, terms that apply only to
birds are noted as such. Most terms that are bolded in the text of the Handbook of Bird Biology appear here. Numbers in brackets
following each entry give the primary pages on which the term is dened.
Please note that this glossary is also available on the Internet at <www.birds.cornell.edu/homestudy>.

abdominal air sacs: A pair of air sacs in the abdominal region

of birds that may have connections into the bones of the pelvis
and femur; their position within the abdominal cavity may shift
during the day to maintain the birds streamlined shape during
digestion and egg laying. [4101]
abducent nerve: The sixth cranial nerve; it stimulates a muscle
of the eyeball and two skeletal muscles that move the nictitating
membrane across the eyeball. [441]
abiotic: Nonliving; includes both things that are dead (such as
dead leaves) and those that have never been alive (for example,
rocks). [97]
accessory nerve: The eleventh cranial nerve; it carries motor
output to constrict the neck muscles. [442]
accommodation: The changes in the curvature of the lens (and
cornea, in birds) of the eye brought about by the action of the
ciliary muscles. These changes allow the eye to focus on objects
at different distances. [450]
acetabulum: At the hip joint, the hollow on the pelvic girdle
into which the head of the femur ts. [424]
acoustic nerve: See vestibulocochlear nerve. [442]
adaptation: A genetically controlled trait that increases an individuals tness relative to that of other individuals. [135]
adaptive: Describes a trait that better promotes an individuals
tness than does some alternative form of that characteristic.
adaptive management: A type of ecosystem management (see
separate entry) in which managers continue to learn more about
the ecosystem as they proceed, and continually modify their
management techniques to incorporate the new information.
adaptive radiation: The evolution, from a common ancestor,
of a variety of different species adapted to different niches; the
species usually have different morphologies and behaviors.
adherent cup nest: A cup nest made of mud or saliva that relies on chemical forces to hold it to a vertical surface; built by
many swifts, including the Edible-nest Swiftlets of Southeast
Asia, whose nests are used in the Asian delicacy birds-nest
soup. [832]
adoption: In avian biology, the peaceful acquisition of a lone
chick or chicks by a pair of adults other than the biological
parents. [8128]
adrenal glands: Small yellow or orange endocrine glands at the
cranial end of each kidney; they produce a variety of hormones
(including adrenaline, steroids, and the sex hormones) that are
involved with circulation, digestion, and reproduction. [474]
advertising displays: Displays performed by one sex (usually
the male) to attract a mate of the opposite sex; also called mate
attraction displays. [637]

aerodynamic valve: A vortex-like movement of air within the air

tubes of each avian lung, at the junction between the mesobronchus and the rst secondary bronchus; it prevents the backow
of air into the mesobronchus by forcing the incoming air along
the mesobronchus and into the posterior air sacs. [4102]
African barbets: A family (Lybiidae, 42 species) of small, colorful, stocky African birds with large, sometimes serrated, beaks;
they dig their nest cavities in trees, earthen banks, or termite
nests. [185]
Afrotropical region: Zoogeographic region including Madagascar, southern Arabia, and all of Africa south of the Sahara
Desert. Sometimes called the Ethiopian Region. [170, 181]
afterfeather: A small feather that grows from the lower shaft of
a contour feather and resembles the main feather but in miniature. [313]
age-specic fecundity: The average birth rate for females in a
particular age group in a population. [964]
age-specic survival rate: The proportion of individuals in a
particular age group in a population that survive a particular
interval of timeusually a year. [964]
age structure: The relative proportions of individuals of different agesusually noted for a given population. [980]
airfoil: Any structure designed to help lift or control a ying
object by using the air currents through which it moves. A typical airfoil, such as the wing of a bird or airplane, is rounded on
top and curved inward below. [510]
air sacs: Thin-walled, transparent sacs extending from the mesobronchi or the lungs to different regions of the body; they act
as bellows to bring air into the body and store it until expiration.
They are found only in birds. [4100]
air speed: A ying individuals speed relative to the air through
which it is moving. It does not include being carried along or
slowed down by the wind, so it may or may not reect a birds
speed relative to the ground. [545]
albino: An individual that lacks the pigment melanin all over
its body. An individual that lacks all types of pigments is called
a complete albino. [352]
albumen: Egg white; albumen is composed primarily of water
and protein. [863]
alimentary canal: The tube for the passage, digestion, and absorption of food; in most birds, it includes the esophagus, crop,
two-part stomach, small intestine, ceca, and the large intestine.
It is also called the gastrointestinal tract, digestive tract, or
gut. [4103]
allantois: In avian biology, the extra-embryonic membrane inside the egg that forms a sac into which the developing embryo
shunts all metabolic wastes that cannot evaporate through the
shell, such as uric acid crystals. [865, 869]
Allee effect: The response, shown in some species when population density falls below some threshold level, by which reproductive behavior and/or social structure become disrupted




in various ways. In some circumstances,

this effect may cause species to be unusually prone to extinction. [1038]
alleles: Alternate forms of genes. Most
animals have two alleles for each trait:
one allele is on the chromosome they received from their mother, and the other is
on the chromosome they received from
their father. As an example consider eye
color in humans, which is determined
primarily by whether someone has two
alleles for blue eyes, two alleles for brown
eyes, or one of each (in which case, the
brown dominates and the person has
brown eyes). [1074]
allopreening: Mutual preening during
which two birds preen each other, usually around the head and neck. In many
species allopreening not only keeps the
plumage clean and orderly, but also
helps to establish social bonds between
individuals. [319]
alpine tundra: Ecosystem found above
the tree line on mountains; it consists of
rugged, well-drained terrain interspersed
with meadows with low-growing vegetation and a profusion of summer-blooming wildowers. Very few birds breed in
this harsh environment. [9114]
alternate plumage: In the HumphreyParkes system of nomenclature, alternate
plumage is the plumage worn by an adult
bird during the breeding season, if that
plumage is produced by a partial molt
before breeding. If a bird does not molt
before breeding, it continues to wear its
basic plumage during breeding. In the
traditional system, the alternate plumage
was known as the nuptial plumage or the
breeding plumage. [333]
altricial: Describes young birds that
hatch undeveloped and in many cases
naked or with sparse down; such helpless young require complete parental
care. [8106]
alula: A group of two to six feathers projecting from the phalanx of the birds rst
nger (its thumb) at the bend of the wing. It
reduces turbulence by allowing ne control of airow over the wing. [111, 515]
American Ornithologists Union (AOU):
The largest organization of professional
ornithologists in North America; it publishes the research journal The Auk and
the Check-list of North American Birds
(commonly called the AOU Check-list;
see separate entry). [139]
amnion: In avian biology, the extra-embryonic membrane inside the egg that becomes lled with uid and surrounds the
developing embryo, allowing it to move
and stay moist, and preventing its various
growing parts from sticking to or blocking
one another. [865]
ampulla: A membranous chamber at the
base of each of the three semicircular

ducts in the inner ear; it contains sensory

hair cells embedded in a gelatinous material and surrounded by endolymph, and
it senses changes in the animals speed or
direction in a particular plane of space.
Information from all three ampullae, one
in each plane, is combined by the brain to
determine the animals motion and thus
to aid balance. [458]
angle of attack: In a ying bird, the angle between the cranial-caudal axis of
the wing and the oncoming airstream.
anisodactyl feet: Foot arrangement in
which the hallux points backward and
the other three toes point forward. Found
in most passerines. [121]
annual survival rate: See survival rate.
ant-acacias: Various species of tropical
and subtropical trees in the genus Acacia that harbor ants inside their hollow
thorns. The ants receive shelter and extra
nutrition from special substances produced by the trees exclusively for the ants,
and in turn keep away insects, mammals,
and other herbivores that might feed on
the trees by attacking them. The ants also
prevent other vegetation from growing
nearby by biting off shoots as they emerge
from the ground. [819]
Antarctic Convergence: Region of the
oceans between about 50 and 60 degrees
south latitude, where cold, north-owing currents meet warmer, south-owing
currents, resulting in large-scale upwelling of nutrient-rich water. The nutrients
support abundant plankton, which attract a great diversity and abundance of
seabirds. [1104]
antbirds: A Neotropical suboscine family
(Thamnophilidae, 197 species) of small,
insectivorous, forest birds; some species
specialize in the technique of following
columns of army ants to prey on the insects and other arthropods stirred up by
the numerous moving ants. [179]
antebrachium: The middle portion of
the forelimb, consisting of the radius and
ulna. The secondary feathers attach to the
ulna. [19]
anterior: Toward the front of an organism, using the earth as a frame of reference. With birds, technically used only
within the eye and inner ear. But in practice, often used interchangeably with the
term cranial. [14]
anterior air sacs: General term referring
to the air sacs nearest the birds front
endthe cervical, clavicular, and anterior thoracic air sacs. [4102]
anterior chamber: Space within the eye
between the iris and the cornea; it is lled
with aqueous uid, which nourishes the
eye and removes wastes. [448]

Cornell Laboratory of Ornithology

anterior lobe of the pituitary gland: Portion of the pituitary gland that receives
instructions from the nervous system in
the form of neurohormones from the hypothalamus. As a result, the anterior lobe
secretes various hormones into the blood
that may act directly on organs or on other endocrine glands (such as the gonads,
adrenals, and thyroids); because of this
central controlling role in the endocrine
system, the anterior lobe is nicknamed
the master gland. [472]
anting, active: Picking up an ant or other
chemically potent object, such as a millipede, and deliberately rubbing it in the
featherspresumably to deter ectoparasites. [322]
anting, passive: Positioning oneself
among a swarm of ants, permitting them
to run all over the body and to move in
and out among the feathers, presumably
to deter ectoparasites. [322]
antiphonal: Describes a singing interaction in which two individuals alternate
their contributions. Often used to describe duets between a male and female
bird. [778]
antorbital fenestra: An opening on each
side of the skull in front of the eye socket;
found in all archosaurs. [E8]
antpittas: Together with antthrushes,
antpittas form the suboscine family Formicariidae (60 species). They are small,
drab Neotropical birds with loud, ringing songs; many haunt the rain forest
oor and may follow army ant swarms.
antthrushes: Together with antpittas,
antthrushes form the suboscine family Formicariidae (60 species). They are
small, drab Neotropical birds with loud,
ringing songs; many haunt the rain forest
oor and may follow army ant swarms.
aorta: The main artery exiting the heart;
its branches distribute oxygenated blood
to all parts of the body. [478]
aortic arch: The curve of the aorta, just
after it exits the left ventricle of the heart.
In birds, the aorta curves to the birds right
as it passes dorsal to the heart and toward
the backbone, but in mammals, the aorta
curves to the left. [483]
AOU: See American Ornithologists
Union. [139]
AOU Check-list: Bird checklist produced
by the Committee on Classication and
Nomenclature of the American Ornithologists Union; it contains common
and scientic names of all birds that occur in North America north of Mexico, or
near North American coasts, including
Hawaii, and is the generally accepted
reference for common names of North
American birds. [164, 1111]

aponeuroses: Shiny, broad sheets of connective tissue that bind muscle bers together to form muscles. [427]
aposematic: Having bright, bold colors
and patterns (often reds and oranges) that
advertise to potential predators that an
individual is bad tasting or poisonous. A
few species that are perfectly palatable
also have evolved these warning colors
and patterns. The strategy of sporting
aposematic coloration is called aposematism. [926]
appeasement displays: Displays given
to decrease the aggression of another
individual; they usually consist of stereotyped postures that de-emphasize
the performers weapons or size and
expose vulnerable parts of its body. For
example, a submissive bird may point its
beak down or away, fold its wings, lower
or turn away its head, point its tail down,
or adopt some combination of these postures. [628]
appendicular skeleton: Portion of the skeleton consisting of the sternum (breast bone),
the pectoral girdle including the front limbs
(wings), and the pelvic girdle including the
hind limbs (legs). [46, 418]
apteria (singular, apterium): Regions of
bare or less-feathered skin between the
feather tracts of birds. [32]
aquatic birds: All birds with webbed
feet that commonly swim, including the
Anatidae; also, all deep-water waders
belonging to the order Ciconiiformes,
such as herons and storks. Also called
water birds. [165]
aqueous uid: A cell-free uid, similar
in structure to blood plasma; it lls the
anterior and posterior chambers of the
eye (between the lens and the cornea),
providing nourishment and waste removal. [447]
arachnoid: The middle of the three vascularized membranes, called meninges,
surrounding the brain and spinal cord.
The meninges provide sustenance and
waste removal for the cells of the brain
and spinal cord, which are not served by
the circulatory system. [436]
arboreal: Living in trees. [E18]
arboreal theory (of the origin of avian
ight): Theory suggesting that ight originated in small, arboreal (tree-dwelling),
reptile-like birds that jumped or glided
among tree branches. Proponents suggest that feathers rst evolved to keep
the animals warmer in the cooler arboreal environment, and then were used to
extend jumps or glides. First proposed
in 1880 by O. C. Marsh. Also called the
trees-down theory. [E18]
Archaeopteryx: A feathered reptile from
150-million-year-old Jurassic limestone
deposits. Because it had a mosaic of

Apo Asi
bills and gaudy, clashing colors. They are
endemic to the Oriental zoogeographic
region. [189]
Asities: Members (with false sunbirds)
of the family Philepittidae, endemic to
Madagascar. These two species of suboscine passerines feed on fruits. [188]
aspect ratio: The ratio of an objects
length to its width. It is used to refer to
the shape of a birds wing: the larger
the aspect ratio, the more elongated the
wing. [537]
asynchronous hatching: Pattern of hatching in which the eggs of a single clutch
hatch over a period of several days, resulting in a brood of young of different
ages. This pattern occurs when incubation begins at the time the rst egg is
laid. Because eggs are laid one per day, at
one- to two-day intervals, the embryos of
the earliest-laid eggs have already started
to develop by the time the later eggs are
laid, and they hatch sooner. [688]
atlas: The rst cervical vertebra; in birds,
it articulates with the single occipital condyle on the base of the skull. (Mammals
have two occipital condyles.) [416]
atria (singular, atrium): The two thinwalled, anterior chambers of the heart;
they receive blood returning to the heart
from the lungs (left atrium) or body (right
atrium). [477]
atrioventricular valve: The valve between
each atrium and its corresponding ventricle; it prevents the backow of blood
into the atrium as the ventricle contracts.
atrophy: To shrivel or die back; may be
pathological or part of the normal course
of development. [427]
attentive periods: In avian biology, periods spent on the nest during incubation.
auditory nerve: See vestibulocochlear
nerve. [442]
auditory tube: Air-lled tube leading
from the middle ear to the throat; it helps
to equalize the air pressure on the two
sides of the eardrum. In birds, the right
and left tubes join and enter the roof of
the mouth caudally through just one
opening. In mammals, the right and left
tubes enter the mouth separately. Also
called eustachian tube. [457]
auricular feathers: A patch of feathers
covering the external ear opening. Their
open texture protects the ear from debris
and wind noise, yet helps to channel
sounds into the ear. [18, 455]
Australasian region: One of the major
zoogeographic regions of the world,
stretching from a line termed Wallaces
Line east of the islands of Timor and Sulawesi in Indonesia, southeast to New
Zealand, and including New Guinea,

Handbook of Bird Biology


bird and reptile characteristics, its relationship to modern birds and other reptiles has been highly controversial since
its discovery in the early 1860s. [E2]
archosaurs: All reptiles in the Archosauromorpha, the large group of diapsid
reptiles that includes all thecodonts and
their descendants (including birds and
crocodiles), but does not include lizards,
snakes, or turtles. All archosaurs have an
opening on each side of the skull, in front
of the eye socket, called the antorbital fenestra. [E8]
arctic tundra: Ecosystem found around
the world in a belt extending north from
the limit of trees; it consists of swampy
atland covered with low-growing vegetation, interrupted by countless shallow
lakes. Very few birds live in the arctic tundra year round, but many migrants breed
there, taking advantage of the long hours
of daylight and numerous spring insects
for feeding their young. [9114]
arctic tundra/coniferous forest ecotone:
Transitional zone between the arctic tundra and coniferous forest ecosystems,
sometimes called the northern timberline. It consists of ngers of open, stunted
spruce forest extending north between
ngers of low, shrubby tundra extending
south. [9123]
area-sensitive species: Species that require large tracts of habitat in order to
persist and breed successfully. [1072]
army ants: A subfamily of primarily
tropical ants that are highly social and
nomadic: colonies make daily raids in
long streams of thousands of individuals
moving across the forest oor, looking for
insects and other arthropods and ushing
out many other small creatures as they
move. Numerous birds (see antbirds) and
some primates follow moving army ant
swarms to take advantage of the prey they
ush. [988]
arterioles: Small blood vessels branching
from arteries. They carry blood from the
arteries to the capillaries. [481, 484]
artery: A vessel conducting blood away
from the heart. All arteries except the pulmonary artery carry oxygen-rich blood.
[481, 482]
articular bone: Bone on the upper surface of each side of the lower jaw of many
vertebrates, near the caudal end; in birds,
it links with the quadrate bone of the upper jaw, forming the joint between the
jaws. [412]
arytenoid cartilages: Two cartilages of
the larynx; they stiffen and hold the shape
of the eshy folds surrounding the glottisthe opening of the larynx. [491]
Asian barbets: A family (Megalaimidae,
26 species) of chunky birds slightly smaller than a Belted Kingsher, with thick


Australia, Hawaii, and other islands of

the mid-Pacic Ocean. Sometimes called
the Australian region. [170, 191]
Australasian robins: A family (Eopsaltriidae, 44 species) of songbirds endemic
to the Australasian region. Reminiscent
of both New and Old World robins in
coloration, they actually are more like
ycatchers, although they usually snatch
food from the ground. [195]
autonomic nervous system: A set of
nerves considered as a group because
of their similar functions. Acting primarily unconsciously, they innervate the
smooth muscle of the viscera, glands,
and blood vessels, thus controlling the
automatic function of the internal organs.
They are under direct chemical control
from substances circulating in the blood.
[443, 444]
avifauna: The set of bird species living in
a region. [169]
axial skeleton: The portion of the skeleton consisting of the skull, hyoid apparatus, and the vertebral column of the
neck, trunk, and tail. [46]
axillaries: A cluster of feathers in the
birds armpit; they are recognizably
longer than those lining the wing. [112,
axis: The second cervical vertebra.
axon: The cable-like, impulse-conducting, main axis of a neuron. [432]


Aus Bio

babblers: A diverse family (Timaliidae,

267 species) of gregarious, insectivorous birds, many of which have complex
social systems and breed cooperatively.
They are found in the Afrotropical, Oriental, and Australasian regions. [190]
back: In birds, the dorsal side of the body,
between the neck and the rump. [16,
barbs: The parallel branches extending
from each side of the rachis of the feather
shaft; collectively they form the vanes.
barbules: Branchlets coming off both
sides of the barbs of a feather, at right angles to the barbs and in the same plane.
Adjacent barbules hook together, holding the vane intact. [33]
basal archosaurs: See thecodonts. [E8,
basal metabolism: The number of calories an organism uses when completely
at rest, which indicates the amount of energy needed to maintain minimal body
functions. [4144]
basic plumage: In the Humphrey-Parkes
system of nomenclature, the plumage

worn by an adult bird for the longest

time each year; it usually is produced
by a complete molt. In the traditional
system, this plumage was known as the
nonbreeding or winter plumage. If a bird
does not molt before breeding, it continues to wear its basic plumage during
breeding. [333]
basilar papilla: The lower membrane
of the cochlear duct in the inner ear of
birds; it is coated with a layer of sensory
hair cells. Sound waves set the basilar papilla into motion, causing the hair cells
to push against the tectorial membrane,
triggering nerve impulses in the hair cells
that are sent to the brain in the process of
sound perception. In mammals, the corresponding structure is called the basilar
membrane. [459]
BBS: See Breeding Bird Survey. [962]
B cells: Special white blood cells found in
the lymphatic tissues; they are important
in the immune response because they
produce antibodies. B cells are produced
by the cloacal bursa and are also important in understanding the development of
AIDS in humans. [4123]
beak: A birds upper and lower jaws, including the external covering; also called
the bill. [16]
bee-eaters: An Old World family (Meropidae, 26 species) of brightly colored
birds with long, slender beaks.They catch
stinging insects in a manner similar to
that of ycatchers, and then beat them to
remove the stingers before eating them.
belly: In birds, part of the lower (ventral)
surface of the body between the breast
and the vent. [16, 19]
bend of the wing: The prominent angle
at the wrist, where the birds wing bends
noticeably. [111]
benets of philopatry hypotheses: A set
of possible explanations for why certain individual birds might forego their
own breeding in a particular breeding
season and act as helpers at the nest of
other breeding pairs (usually their parents or other close relatives); the benets
of philopatry hypotheses focus on the
possible benets to young adults of remaining with their parents. Examples of
these hypotheses include (1) the survival
of young adults may be improved when
they remain in a group, (2) by helping,
young adults may improve the survival
of close relatives (and thus increase their
own indirect tness), and (3) by staying
with their parents, young adults may increase their own chance of acquiring a
superior territory, either by monitoring
vacancies in neighboring sites or by inheriting their natal territory. [689]
Bergmanns rule: Rule describing the
pattern of body sizes found within most

Cornell Laboratory of Ornithology

bird and mammal species, in that individuals living in colder regions tend to be
larger than those living in warmer areas.
Bernoullis law: Physical law stating that,
in any system of airow, static and dynamic pressure must always add up to a
constant. Because the airow over a moving airfoil (such as a birds wing) is faster
above than below, the dynamic pressure
is higher and thus the static pressure is
lower. Because the static pressure below
the wing is higher than that above, lift
is created and the bird can remain aloft.
biconical: Describes an egg that is slightly longer than subelliptical; also called
fusiform or long subelliptical. [873]
bile: Substance, produced by the liver,
that emulsies fats to facilitate their digestion. In birds with no gall bladder, bile
is released directly into the small intestine
through the hepatoenteric ducts; in birds
with a gall bladder (and mammals), bile
is stored in the gall bladder and released
through the bile ducts. [4124]
bill: A birds upper and lower jaws, including the external covering; also called
the beak. [16]
bill tip organ: An aggregation of sensory
cells at the tip of both the upper and lower
beak, best developed in ducks, geese,
sandpipers, and snipe; it is thought to
sense tactile stimuli during feeding.
bill-wiping: A maintenance behavior in
which a bird swipes its bill sideways on
tree branches, the ground, or other surfaces, especially after eating messy foods
such as oily insects or suet. [340]
binocular vision: A type of vision that
produces three-dimensional images,
in contrast to monocular vision, which
produces at images. Binocular vision
results when the eyes are positioned toward the front of the head, so that objects
are detected by both eyes simultaneously.
binomial nomenclature: The currently
accepted system of naming organisms,
devised by Linnaeus, in which each species is designated by two words: the genus and the species names. [147]
bioaccumulation: The increase in the
concentration of toxic or other foreign
substances in organisms bodies as a result
of taking up the substances from the environment (through plant roots, or in ingested
food or water) at a rate higher than that at
which they are excreted from the body.
Many toxic substances that do not occur
naturally, such as DDT and PCBs, are readily deposited in body tissues but excreted
at very slow rates, because organisms have
not evolved mechanisms to metabolize
them effectively. [9126, 9127]

bioconcentration: The stepwise increase,
found at each higher level of the food
chain, in the concentration of certain
chemicals in the bodies of organisms.
Chemicals that usually bioconcentrate in
the food chain are those toxins, such as
DDT, that tend to accumulate (see bioaccumulation) in organisms because they
are taken up from the environment faster
than they are excreted. Also called biological magnication. [9125, 9127]
biodiversity: The great wealth of living
organisms that occur on earth. [1106]
biogeography: The study of the distribution patterns of living things. [950]
biological indicators: See indicator species. [10108]
biological magnication: See bioconcentration. [9126, 9127]
biomass: The total mass of all the living
organisms in a particular population,
community, or area at a given point in
time. [987]
biome: A major terrestrial ecosystem;
in North America north of Mexico, the
nine biomes are tundra, coniferous forest, deciduous forest, grassland, southwestern oak woodland, pinyon-juniper
woodland, chaparral, sagebrush, and
scrub desert. Biomes are named for the
dominant vegetation of the regions climax community, but they include all
the living organisms in that community,
the physical surroundings, and all the
ecological processes that occur there.
biotic: Living. [97]
bipedal: Walking (or running) on two
legs. [E16]
bird community: See community.
birds-of-paradise: A family (Paradisaeidae, 46 species) of forest-dwelling songbirds found primarily in New Guinea;
they are famous for the spectacular, colorful plumages and displays of the males.
birth rate: The number of young born to
an individual or set of individuals, or born
into a population or species, in a given
period of time (often a year). Usually
(especially for humans) expressed as the
number of births per 1,000 individuals
per year. Also called fecundity. [949]
blastoderm: In avian biology, the attened
disc of dividing cells that lies on the upper
surface of the yolk and is the rst stage in
the development of the embryo. [863]
blind: A structure used to conceal a person so that he or she may observe birds or
other wildlife; known as a hide in Great
Britain. [229]
blind spot: Site on the retina where the optic nerve penetrates the retina and leaves

Bio Bro
the eye; because no rods or cones are
present at this spot to capture incoming
light, an object whose image falls on the
spot is not perceived. [449]
blocking: A method of estimating the size
of large ocks of birds by counting the
birds in a block of typical density, beginning at the trailing end of the ock (so
that birds are not ying into the area you
are counting), and then visually superimposing the block onto the rest of the ock
to see how many times it will t. [256]
blood/brain barrier: The specialized arrangement of capillaries in the brain that
prevents blood from reaching the nerve
cells, thus protecting the brain from potentially toxic substances that circulate
in the blood. Capillaries penetrate the
meninges surrounding the brain, but do
not reach the actual neurons of the brain.
blood plasma: See plasma. [486]
blood vascular system: Alternate name
for the circulatory system. [476]
body downs: The down feathers of adult
birds, found under the contour feathers.
Body downs are most common in water
birds and hawks, as they provide extra
insulation. [316]
bone: Tissue composed of living cells in
a mineralized matrix; the bones provide
support for the body and attachment sites
for the muscles. [43]
bony labyrinth: The bony, outer system of
uid-lled canals (containing perilymph)
that make up the inner ear. It forms the
cochlea, semicircular canals, and vestibule, and encloses the membranous
labyrinth. [457]
booming sacs: Brightly colored outpocketings of the esophagus of some North
American grouse that appear at the sides
of the neck; they ll with air and act as
resonators to produce loud sounds during displays. [4114]
booted podotheca: A smooth podotheca,
divided into long, continuous, nonoverlapping scales. [120]
boreal forest: Coniferous forest ecosystem dominated by spruce and r trees
and found around the world, generally
in a belt north of temperate zone deciduous forests and south of the arctic
tundra. Many birds migrate to the boreal
forests to breed, taking advantage of the
long daylight hours and abundant insects
for feeding their young. Some bird species are year-round residents. Also called
taiga. [9116]
bounding: A ight pattern in which a bird
alternates apping (during which it rises
slightly) with glides on closed wings (during which it descends slightly). [526]
bower: A complex mating structure built
by male bowerbirdsmembers of the

Handbook of Bird Biology


passerine family Ptilonorhynchidae. See

bowerbirds. [676]
bowerbirds: A passerine (oscine) family
(Ptilonorhynchidae, 20 species) of New
Guinea and Australia whose polygynous
males attract females by building and
decorating remarkably complex bowers out of twigs and other objects. See
bower. [676]
brachial plexus: Plexus along the spinal
cord of birds at the level of the wing; it is
associated with a cervical enlargement
of the spinal cord. [439]
brachium: The upper (proximal) portion
of the forelimb (wing); it contains the humerus. [19]
brain stem: See medulla oblongata.
breast: In birds, part of the lower (ventral)
surface of the body, between the throat
and belly. [16]
Breeding Bird Survey (BBS): A count
of the breeding birds of North America
conducted each summer since 1967 and
coordinated by the U. S. Fish and Wildlife Service. Observers count birds seen
or heard during three-minute periods at
half-mile (0.8-km) intervals along a 25mile (40-km) stretch of road. Because
the same routes and stops are sampled
each year, BBS data can be used to track
population trends, and the results can be
correlated with habitat. [962]
breeding plumage: See alternate plumage. [333]
breeding season: The period of time during the year when a particular species
may breed. [811]
bristles: Highly specialized contour
feathers in which the rachis is stiffened
and lacks barbs along its outermost parts.
broadbills: A family (Eurylaimidae, 15
species) of stocky, brightly colored, suboscine birds found throughout forests
and scrublands of the Old World tropics, particularly in the Oriental region.
Broadbills use their wide, at, colorful
bills to snatch up large insects. [181]
bronchi (singular, bronchus): The two
major air tubes branching off the lower
end of the trachea; one goes into each
lung. [492]
brooding: Sitting on hatched young, or
sheltering them under the wings, primarily to keep them warm, but also to protect
them from sun, rain, or predators. Occurs
either in the nest, or outside the nest in
those species whose young leave the nest
shortly after hatching. [895]
brood parasite: A bird that lays eggs in
the nests of other species, leaving the
resulting young to be raised entirely by
the host parents. Some species parasitize


others only occasionally; others, called

obligate brood parasites, never build
their own nests, and lay all their eggs in
other species nests. [8139]
brood patch: A patch of skin on the breast
and belly of birds that has lost feathers
and become swollen through both the
retention of large amounts of water in the
tissues and the expansion of blood vessels feeding the skin. It develops a few
days before egg laying in most individual
birds (either male or female) that incubate their eggs by sitting on them, and
increases the efciency of heat transfer
to the eggs. One large patch or several
smaller ones may develop, depending on
the species. [794]
brood reduction: Practice carried out by
some parent birds whose young hatch
asynchronously and thus vary in size,
in which the parents rst feed the most
vigorously begging offspring until it can
swallow no more, and then move on to
another. Thus, in years with low food supplies only the largest and strongest young
will survive, but in years with abundant
food smaller young will survive as well.
Brood reduction ensures that at least
some young will survive in years when
food is scarce. [898]
Brown-headed Cowbird: See cowbird.
[8142, 8148]
bursa of Fabricius: Former name for the
cloacal bursa. [4123]
bustards: A family (Otididae, 25 species)
of large, heavy-bodied, at-headed birds
with long legs and necks. These grounddwelling birds are strong runners, and
frequent open areas in the Oriental,
Australasian, and southern Palearctic
regions, although most species live in
Africa. [183]


Bro Cen

calamus: The hollow lower portion of the

feather shaft, part of which lies beneath
the skin; it has no vanes. [33]
call notes: Bird sounds that are generally
shorter and simpler than songs. Many
seem to convey a specic message, such
as begging calls (hunger), alarm calls
(danger), and contact calls (the callers
location). [714, 772]
CAM: See chorioallantoic membrane.
camouage: Coloration of an organism or
structure serving to conceal it from predators, other enemies, or prey. [361]
canopy: The upper, continuous level
of vegetation in a forest; it contains the
branches of tall, mature trees and the epiphytes that grow on these branches. In
dense forests, the canopy receives most
of the sunlight and thus has the greatest

productivity. The tallest trees that stick

above the main canopy, somewhat like
lollipops, are not considered part of
the canopy; they are called the emergent
layer. [993]
capillaries: The smallest blood vessels.
Within all body tissues except the epidermis and those in the central nervous
system, capillaries form networks called
capillary beds, which connect the arterioles and venules. The thin walls of
the capillaries allow materials to be exchanged between the blood and the body
cellsthe body cells absorb oxygen from
the blood, and the blood takes up wastes
from the body cells. [481]
capillary bed: See capillaries. [481]
cardiac muscle: A special type of smooth
muscle that forms the bulk of the heart.
Cardiac muscle can contract without
being stimulated by the nervous system.
carina: See keel. [423]
carinates: Birds that have a keel (carina)
on their sternum; includes all ying birds
and many smaller ightless ones. [423]
carnivores: Meat-eaters. [9123]
carotenoids: Pigments producing bright
yellow, orange, or red colors. They are
synthesized only by plants, so birds must
obtain them via their diets. [351]
carotid artery: Main artery supplying the
head and neck region; among birds it is
highly variable, with different species
having none, one, two equal in size, or
one large and one small. [484]
carpals: The bones of the wrist. In birds,
they are fused and reduced to just two
bonesthe radiale and ulnare. [19]
carpometacarpus: The largest bone of
the manus of birds, formed by the fusion
of some of the carpals (wrist bones) with
the metacarpals (palm bones). [19]
carrion: The esh of dead animals.
carrying capacity: The maximum population size or density that a particular area
can support over the long term, without
any degradation in the quality of the area
or its resources. [970]
cartilage: A tissue with living cells embedded in a nonmineralized matrix; cartilage is found in the exible joints and is
capable of growth or resorption as well as
transformation into bone. [46]
casques: Enlargements on the top of the bill
or the front of the head, usually involving
the underlying bone. Found in cassowaries
and hornbills, among others. [348]
cassowaries: A family (Casuariidae, 3
species) of large, ightless ratites inhabiting New Guinea and Australia. They have
a distinctive bony casque on top of their
head. [349]

Cornell Laboratory of Ornithology

cast: See pellet. [4119, 4120]

caudal: 1. Toward the tail or posterior part
of the body. [14] 2. Pertaining to the
tail. [415]
caudal mesenteric vein: Vein that in birds
brings blood from the lower portion of
the digestive tract to the renal portal veins
that form the venous ring connecting the
lobes of the two kidneys; it is part of the
avian renal portal system. [485]
caudal vena cava: Single large vein that
gathers blood from veins coming from
the legs, tail, kidneys, and caudal regions
of the body and delivers the blood to the
right atrium of the heart. [483, 484]
caudal vertebrae: The vertebrae of the
tail. In birds, the anterior caudal vertebrae fuse with the sacral and lumbar vertebrae and some of the thoracic vertebrae
to form the synsacrum; the next four to
nine vertebrae articulate freely; and the
posterior ones fuse to form the pygostyle
(tail bone). [415, 418]
caval veins: General term for the right
and left cranial vena cavae and the caudal vena cava; these large veins gather
deoxygenated blood from the body and
return it to the right atrium of the heart.
[477, 484]
cavity adopters: Birds that nest in cavities
but do not excavate their own, instead
obtaining cavities that were created by
physical forces (such as decay or erosion)
or by other species. Bluebirds, Great
Crested Flycatchers, Tree Swallows, and
House Wrens are all cavity adopters. Also
called secondary cavity nesters. [839]
CBC: See Christmas Bird Count. [960]
cell: Membrane-enclosed unit capable of
metabolism and reproduction; the basic
structural and functional unit of life. [42]
cell body: The part of a neuron containing
the nucleus and surrounding cytoplasm;
also called a soma. [432]
center of origin (of a species): The geographic area where a species evolved.
central fovea: Area in the central part of
the retina of most birds and mammals
where the cones are most concentrated
and the neural layer (the nerves from the
rods and cones, which overlay the rods
and cones and block some light) is the
thinnest, and thus vision is the sharpest.
The central foveae provide sharp monocular views of the areas to the sides of
the bird. [449, 453]
central nervous system (CNS): The brain
and spinal cord. [435]
centrum: The main body, or central axis,
of a vertebra; the anterior end of the centrum connects to the preceding vertebra,
and the posterior end connects to the following vertebra. [415]

cere: A leathery band of skin covering
the base of the bill, into which the nostrils open; presumably the cere protects
the nostrils. Present only in certain birds,
such as hawks, pigeons, and some parrots. [340]
cerebellum: A large, deeply folded
structure on the dorsal surface of the
hindbrain, attached to the brain stem by
two pairs of stout neural tracts; it controls muscular coordination and plays an
important role in balance, posture, and
proprioception. The cerebellum of birds
is particularly large, due to the demands
of ight. [437]
cerebral hemispheres: The two large,
smooth lobes on the dorsal anterior region
of the forebrain; together they form the cerebrum, which coordinates and controls
complex behaviors, including memory
and learning. In mammals, these lobes
have folds and grooves. [436]
cerebrum: See cerebral hemispheres.
cervical air sacs: A pair of air sacsone
sac on each side of the bodylocated in
the neck region of birds; one sac usually
extends from each lung, but sometimes
a series of cervical sacs are located along
the neck, as in geese. [4101]
cervical enlargement: A swelling along
the spinal cord at the level of the wings,
associated with the brachial plexus.
cervical vertebrae: The vertebrae of
the neck region; the cervical vertebrae
of birds have uniquely shaped centrum
ends (see heterocoelous centrum ends).
chalazae: In avian biology, the gelatinous, usually milky white, stringy coils
of albumen (egg white) that surround and
protect the egg yolk, and are visible at either end of the yolk as twisted cords. The
chalazae attach to the far ends of the eggshell and form a suspension system for
the yolk that allows it to rotate throughout
embryonic development. [862]
channelization: The process by which
humans deepen and straighten natural
streams, converting them into waterlled ditches. Theoretically channelization controls ooding along the stream,
but it destroys the stream ecosystem and
often increases ooding downstream.
chaparral: North American ecosystem
found on the low hillsides of southwestern California; it consists of dense stands
of broad-leaved evergreen shrubs, dominated by chamise and manzanita. This
ecosystem is dry, with long hot summers
with frequent res; rain falls only in winter. A moderate number of birds breed in
the chaparral. [9120]

Cer Cla
checklist: A printed list of the birds found
in a particular area. [246]
cheek: See malar region. [17, 18]
chin: A small area under the lower beak
of birds. [17, 18]
choana: A single slit in the roof of the
mouth, running in an anterior-posterior
direction, through which the two nasal
cavities open to the mouth. [490]
chorioallantoic membrane (CAM):
Membrane covering the entire inner
surface of the avian eggshell, inside the
inner shell membrane; it develops partway through embryonic development
from a fusion of two extra-embryonic
membranes, the chorion and allantois.
The CAM is richly invested with blood
capillaries, and together with the pores in
the eggshell, it allows the embryo to carry
out gas exchange as it receives oxygen
from outside the egg and expels carbon
dioxide. [866, 869]
chorion: In avian biology, the extra-embryonic membrane inside the egg that surrounds the entire avian embryo and the
other three extra-embryonic membranes
(the allantois, amnion, and yolk sac). It is
homologous (evolutionarily related) to
the mammalian membrane, also called
the chorion, which forms much of the placenta in most mammals. [865, 869]
choroid: The middle layer of the three
main layers of the eye; it lies just inside
the sclera. It is pigmented and forms the
iris and ciliary processes. [448]
Christmas Bird Count (CBC): A count
of the wintering birds of North America
conducted each year since 1900 and
coordinated by the National Audubon
Society. Observers count as many individual birds as possible within one of the
circular count areas 15 miles (24 km) in
diameter that are scattered across North
America and beyond. Observers record
their time spent and distance covered,
so the numbers of birds seen can be adjusted for observer effort. The data can be
used to track winter bird distribution and
abundance, as well as long-term population trends. [960]
chromosomes: Long strands of DNA
found in the nuclei of most cells. Each
section of the chromosome (a specic
sequence of nucleotides) that codes for
a specic protein is called a gene. [144,
ciliary muscles: Muscles that, in birds, attach to the ciliary processes, which attach
to the lens of the eye. When the ciliary
muscles contract, they move the ciliary
processes, which squeeze the lens and
make it become more round. In mammals, ciliary muscle contraction relaxes
the lens, allowing it to become round by
elastic rebound. [448]

Handbook of Bird Biology


ciliary processes: Structures of the choroid of the eye that attach to the lens and
hold it in place. Ciliary muscles move the
ciliary processes, which in turn move the
lens, changing its shape. Only in birds do
the processes attach directly to the lens.
circadian rhythms: Daily cycles of behavioral and physiological events exhibited
by organisms; they are regulated by an
internal biological clock and persist even
when organisms are kept under constant
environmental conditions. Examples include daily patterns of activity, body temperature, or nectar production. [561]
circannual rhythms: Cycles of behavior,
growth, or other physiological activities
that occur on approximately a yearly
basis; like circadian rhythms, they are
regulated by an internal biological clock
and persist even when organisms are
kept under constant environmental conditions. Examples include yearly patterns
of migration, shedding and regrowth of
antlers, and hibernation. [562]
circumpolar constellations: The stars
near to and surrounding the North Star
(Polaris). [587]
cisticolas: A family (Cisticolidae, 117
species) of small, drab, insectivorous
warbler-like birds of open, grassy areas
of the Old World, primarily Africa. [87]
CITES: See Convention on International
Trade in Endangered Species. [1093]
class: Level of classication of organisms above order and below phylum; similar orders are placed within
the same class. All birds are in the class
Aves. [152]
clavicle: The collar bone; in nearly all
birds, the left and right clavicles fuse with
a small interclavicle bone to form the Vshaped furcula (wishbone). [419]
clavicular air sac: A single, median air
sac between the clavicles and surrounding the bifurcation of the trachea of birds.
claw arc: The angle between the tip and
base of a claw, considering the claw as a
section (arc) of a circle; used by researchers to compare the curvatures of different
claws. It is signicant because claw curvature is related to a birds habits: ground
dwellers have atter, straighter claws
(smaller claw arcs), perching birds have
moderately curved claws, and trunk
climbers have the most highly curved
claws (largest claw arcs). [E6]
clay lick: Clay banks rich in minerals,
such as calcium, that are visited by frugivorous or seed-eating birds, such as
parrots and macaws, who eat the clay to
obtain minerals that are otherwise lacking in their diet. Clay licks are one type
of mineral lick. [928]



Cle Con

Clean Water Act: Federal law passed in

the United States in 1972 that attempted
to improve the quality of surface waters
by controlling pollution and requiring
sewage treatment. Section 404 (see separate entry) contains a set of protections
for wetlands. [1093]
climax community: The association of
plants and animals (and other organisms)
that can perpetuate itself in a given area
in the absence of large-scale climatic
change, disease, or human disturbance.
The climax community is the nal stage
in the process of ecological succession in
any given area. [9109, 9110]
cline: A gradual change in certain characteristics of individuals of the same species, which is evident in a geographic
progression from one population to the
next. [155]
cloaca: Common opening at the lower end
of the avian digestive tract for the digestive, excretory, and reproductive systems;
it receives feces from the large intestine,
uric acid from the kidneys, and eggs or
sperm from the gonads, and releases these
materials through the vent. [4123]
cloacal bursa: A lymphoid organ that
opens into the roof of the cloaca in young
birds and atrophies in later life; it produces special white blood cells called
B cells, which are important in immune
function and are of interest to researchers
for their role in the development of AIDS
in humans. Previously known as the bursa of Fabricius. [4123]
cloacal phallus: The copulatory organ of
male ratites and waterfowl; it is an elongate, spiral, ridged structure that erects
by lymphatic pressure during copulation.
Sperm travel along its surface to reach the
cloaca of the female. [4129]
cloacal protuberance: A swelling at the
caudal end of the deferent ducts, visible externally in hand-held birds with
the feathers parted; it is present only in
breeding male passerines, and is used by
researchers to determine sex and breeding condition. The swelling is caused
by the enlargement of structures in the
terminal regions of each deferent duct
during the breeding season, and may
help to keep sperm slightly cooler than
the bodys core temperature (as does the
scrotum of mammals). [4129]
clutch: A complete set of eggs; those laid
in an uninterrupted series, for a single
nesting, by one female. [878]
clutch size: The number of eggs in a given
clutch. [878]
CNS: Central nervous system; the brain
and spinal cord. [435]
coastal: Of the coast; coastal bird species primarily occupy the shallower waters around oceanic islands or above the

continental shelf, feeding mainly on sh,

crustacea, and mollusks, which they nd
on or near beaches and other shorelines.
They visit land frequently, during both
the breeding and the nonbreeding season. [198]
cochlea: Elongate, bony, structure of the
inner ear that is concerned with hearing;
it is part of the bony labyrinth and thus is
lled with perilymph. It contains the perilymph-lled vestibular and tympanic canals, which are connected to one another
at one end, and the endolymph-lled cochlear duct, which lies between them. In
mammals, but not in birds, the cochlea is
coiled like a snails shell. [459]
cochlear duct: Membranous canal
inside the cochlea, bounded above by
the tectorial membrane and below by
a membrane that in birds is called the
basilar papilla. The cochlear duct is part
of the membranous labyrinth and thus is
lled with endolymph. [459]
cochlear window: A soft spot at the end
of the bony cochlea farthest from the
vestibular window; it acts both as a pressure-release valve and as a damper for the
waves in the cochlea. Formerly called the
round window. [459]
coevolution: An evolutionary interaction
between two or more species in which
one evolves an adaptation that affects
another, and then the other evolves an
adaptation in response, and then the rst
evolves another adaptation in response
to the response, and so on. Coevolution
sometimes results in a kind of battle or
arms race between two species. Notable examples of coevolution include
interactions between predators and prey,
plants and pollinators, and brood parasites and hosts. [8146]
cohort: A group of individuals born during
the same time period (often a year). [85]
cold-blooded: See ectothermic. [11]
cold front: The interface between a mass
of cold air and the warm air mass it is overtaking; the dense, cold air tends to wedge
under the warm air, forcing the warm air
up and cooling it abruptly, forming precipitation that is often accompanied by
strong winds and lightning. [569]
colic ceca (singular, cecum): Pouches
extending from the junction between
the small and large intestines that hold
partly digested food long enough for
bacterial action to further break it down;
digested material is released to the large
intestine, where any released nutrients
are absorbed. Birds may have none, one,
or one or two pairs of colic ceca, and
the size is highly variable among species.
Also spelled caeca or caecum. [4123]
colies: See mousebirds. [185]
color phases: Polymorphisms in which the

Cornell Laboratory of Ornithology

morphs differ in color; for example, the

red and gray phases of Eastern ScreechOwls and Ruffed Grouse, and the blue
and white phases of Snow Geese. [981]
columella: Small, thin bone extending
across the middle ear of birds, attached
at one end to the inner surface of the
eardrum and at the other end to the vestibular window of the inner ear; it transmits sound waves from the eardrum to the
uid-lled cochlea. [456]
comb: Fleshy, erect structure positioned
longitudinally on top of the head of a
bird, often with a serrated margin (like
a hair comb), as in domestic chickens.
commensal: Describes a relationship
between two species or individuals in
which one benets and the other neither
benets nor is harmed. For example,
when Cattle Egrets follow large herbivorous mammals to eat the insects they stir
up as they move, the egrets benet but
the mammals appear unaffected in any
way. [991]
communal roost: A group of birds gathered to spend the night together, sleeping;
may consist of just one species or a number of different species. Birds that form
particularly large and noisy communal
roosts include vultures, ravens, crows,
starlings, herons, egrets, ibis, grackles,
blackbirds, cowbirds, robins, and the
extinct Passenger Pigeon. [665]
community: All the populations of species living and interacting with one another in the same place. Communities
also can be dened as containing only
certain types of species in one location,
such as a forest bird community or a
stream insect community. [93]
competitive exclusion principal: Rule
stating that no two species can occupy
exactly the same ecological niche in a
community; if they did, then eventually
one would outcompete the other and
cause it to go extinct. [9102]
complete albino: An individual lacking
all types of pigments in the plumage,
eyes, and skin. [352]
complete molt: A type of molt in which
the entire feather coat is replaced.
conchae: Two thin, scroll-like structures
(one median, one anterior) extending
from the lateral wall of each nasal cavity;
they are covered with a mucus-secreting
membrane that contains the nerve endings of the olfactory nerves, which sense
odors. The mucus traps dust, and blood
vessels in the membrane warm the inhaled air. [490]
cones: One of the two kinds of light-sensitive cells lining the retina of the eye;
they are responsible for visual acuity (due

to their tight packing) and for sensing color information (via four or ve pigments
and specialized oil droplets in birds), but
are not very sensitive to low light levels.
When light energy stimulates a cone cell,
it sends a nerve impulse to the brain via
the optic tract. [448]
conical: See pyriform. [873]
coniferous forest: Ecosystem of colder
parts of the temperate zone, where there
is sufcient moisture to support a forest;
cone-bearing trees, especially spruce
and r, dominate the vegetation. North
America contains three main types of
coniferous forests: western montane
forest, boreal forest (taiga), and eastern
montane forest. See each forest type for
more information. [9116]
coniferous forest/deciduous forest
ecotone: Transitional zone between the
coniferous forest and deciduous forest
ecosystems; it has a mixture of the two
types of trees and, in North America, has
more species of breeding birds than any
other region. [9123]
conservation biology: An applied science that combines information gained
through the biological elds of ecology,
population biology, animal behavior,
and genetics to attempt to reverse the
widespread declines and extinctions of
species occurring throughout the world
today. [1039]
conservation plan: See habitat conservation plan. [1092]
Conservation Reserve Program (CRP):
Part of the 1985 Food Security Act that
authorized the USDA to lease millions of
areas of marginal croplands from farmers
each year, paying them to keep the land in
perennial vegetation to reduce soil erosion and crop surpluses, and to restore
wildlife habitat. The program has helped
prairie songbirds by providing millions of
acres of grassland habitat. [1035]
conspecics: Members of the same species. [366]
contact call: A sound produced by a
bird that appears to inform a nearby bird
(usually a family member) of the callers
location. Often uttered by a mated male
and female as they forage relatively close
to one another. [773]
contour feathers: Feathers that make up
the exterior surface of a bird, including
the wings and tail; they streamline and
shape the bird, and usually have well-developed barbules and hooklets. [35]
Convention on International Trade in Endangered Species (CITES): International
agreement to which 150 nations voluntarily subscribe that binds participating
parties to monitor, regulate, or prohibit
the import and export of species that the
group has deemed worthy of global pro-

Con Cou
tection. The species are listed in three Appendices: Appendix I lists the most endangered species, for which all commercial
trade is prohibited, Appendix II lists species that would be in immediate danger if
trade were not regulated, and Appendix III
lists species added by individual countries
that are requesting international help in
regulating their trade. [1093]
convergent evolution: The process by
which organisms evolve similar forms,
behaviors, or ecological characteristics
not because they are related, but to meet
similar environmental challenges. [140,
cooperative breeding: Breeding system
in which adults other than the breeding
pair help the breeding pair to rear their
offspring. In birds, if the helpers assist
with incubation or care of nestlings, they
are called helpers at the nest (see separate entry). [688]
cooperative foraging: A technique in
which a group of individuals work together to obtain prey. For example,
American White Pelicans may swim in a
line or semicircle and beat their wings to
drive schools of sh into shallow water.
Other pelicans, cormorants, and mergansers, among others, forage cooperatively some of the time. Also called social
foraging. [664]
coracoids: Strong, stout, paired bones of
a birds pectoral girdle; they are not present in mammals. During ight the coracoids function as a powerful brace holding the shoulder joint, and thus the wing,
away from the body while the pectoral
muscles pull on the wing in the opposite
direction. [419]
corcoracids: The two members of the passerine (oscine) family Corcoracidaethe
blackbird-like White-winged Chough
and the smaller, seed-eating Apostlebird. These large, Australian cooperative
breeders range over agricultural elds in
huge ocks when not breeding. [195]
core area: The portion of a large, continuous habitat, such as a forest, that is far
from the edges and thus is suitable to host
species that would be adversely affected
if forced to live near the edges. [9100]
cornea: The transparent anterior surface
of the sclera; it allows light to enter the
eye. [448]
coronary arteries: Arteries (usually two)
arising from the rst part of the aorta; they
carry blood that nourishes and supplies
oxygen to the heart muscle itself. [479]
coronary veins: Veins that carry deoxygenated blood and wastes from the muscle tissue forming the heart back into the
right atrium of the heart. [479]
corpuscles: An alternate name for blood
cells. [486]

Handbook of Bird Biology


corridors: In conservation biology, long,

narrow areas of wildlife habitat that connect larger areas, thus allowing individuals to move between the larger areas.
countercurrent exchange system: A system in which two uids (liquids or gases)
ow adjacent to one another, but in opposite directions, while heat energy or
materials move passively from one to the
other, from higher to lower temperature
or from higher to lower concentration.
An example is the countercurrent heatexchange system located in the legs of
many gulls and waterfowl: warm blood
on its way to the feet ows through a network of small vessels, which intertwines
with another network of small vessels
carrying cold blood owing from the feet
back to the body core. As a result, the
blood returning from the feet is warmed,
conserving body heat. [4151] Another
example is the countercurrent exchange
system that efciently accomplishes
gas exchange in a birds lungs: blood in
the capillaries around the parabronchi
moves in the opposite direction to the air
in the air spaces. [4100]
countershading: A type of coloration in
which an organism is darker on top than
below; countershading provides camouage by reducing the contrast between
the top and shadowed underside of an
organism so that it appears less three-dimensional. [363]
countersinging: Interaction in which two
birds sing back and forth to each other,
alternating their songs. [736]
coursers: Together with pratincoles, form
the family Glareolidae (17 species). These
slender, plover-like ground nesters live in
open areas in tropical parts of the Old
World, and sometimes cool their eggs or
young by partially burying them in sand
or by bringing water to the nest in their
breast feathers. [183]
courting nests: Nests, usually unlined,
produced by male Winter Wrens and
some male Marsh Wrens to attract females. A male may build a number of
courting nests on his territory. In Marsh
Wrens, females may add a lining and lay
eggs in the courting nest of their chosen
mate. [824]
courtship displays: Displays performed
for the opposite sex to acquire a mate of
the same species, maintain a pair bond,
and/or stimulate and synchronize breeding behavior. [634]
courtship feeding: Feeding of a female
by her mate that occurs during courtship and incubation, often in response to
ritualized begging calls and postures that
resemble those of the young. The function of courtship feeding is unknown, but
hypotheses (which are not mutually ex-




clusive) include (1) it strengthens the pair

bond, (2) the food improves the females
condition, and (3) it allows the female to
assess the males ability to provide food
for the young. [66, 8104]
coverts: The smaller feathers that partly
overlie the ight feathers of the wing and
tail at their bases, like evenly spaced
shingles on a roof. For more information,
see specic types. [111, 113]
cowbirds: Six species of dark, slender,
medium-sized birds that forage on seeds
and insects; they are members of the New
World family Icteridae (blackbirds and
New World orioles). All species except the
Bay-winged Cowbird are obligate brood
parasites: they lay their eggs only in the
nests of other species. The Brown-headed
Cowbird, found throughout the United
States and much of Canada, successfully
parasitizes more than 140 host species and
is thought to have caused severe population
declines in some. It is the only brood parasite common in North America north of
Mexico. [8142, 8148]
cracticids: Members of a distinctive family (Cracticidae, 10 species) of songbirds
endemic to Australasia. They have stout,
straight beaks; loud, melodic calls; and a
generalist, crow-like diet of small vertebrates, eggs, insects, and fruits. Cracticids include butcherbirds, currawongs,
and the Australasian Magpie. [195]
cranial: 1. Toward the head. In practice,
may be used interchangeably with anterior (but see separate entry). [14] 2.
Pertaining to the head, brain, or skull.
cranial kinesis: The ability of the birds
upper jaw (upper beak) to move upward
at the same time that the lower jaw (lower
beak) is depressed, an action permitted
by the highly exible craniofacial hinge.
cranial nerves: Twelve sets of paired
nerves, each with a specic function,
serving the head, neck, and thorax region.
Most exit from the medulla oblongata of
the brain. [437]
cranial vena cavae: Two large veins (right
and left) that gather deoxygenated blood
from large veins coming from the wings,
head, and neck and deliver it to the right
atrium of the heart. [483, 484]
craniofacial hinge: Flexible joint where
the upper beak connects to the rest of the
skull. Also called the nasal-frontal hinge.
cranium: The part of the skull enclosing
the brain; the braincase. [411]
creche: An assemblage of the still-dependent young of two or more (usually
many) breeding females, attended by one
or more adults. Bird species whose young
form creches include some pelicans, a-

mingos, geese, penguins, parrots, jays,

and terns; creches of King Penguins and
Emperor Penguins may contain several
thousand young. Creche formation allows some birds (notably penguins) to
conserve energy. It also may decrease
predation on the young, and may free
parents to spend time foraging in distant
areas, thereby allowing them to bring
more food back for the young and to prepare themselves for the next breeding.
crepuscular: Active at dawn and dusk.
crest: Tuft of feathers on the peak of the
head that either stick up or can be raised.
cricoid cartilages: Two major cartilages
that make up the sides and oor of the
larynx. Also called the laryngeal cartilages. [491]
crop: A dilation of the lower esophagus
that stores food; it is found in many birds
that eat dry seeds or fruit containing
seeds. [4115]
crop milk: Milk-like substance produced
by pigeons and doves; it is composed of
uid-lled cells sloughed from the lining
of the crop and is regurgitated to feed to
nestlings. Crop milk is high in lipids and
vitamins A and B, and has a greater protein and fat content than human or cow
milk. Also called pigeons milk. [4116]
cross-fostering experiments: Studies in
which young of one species are placed
with host parents of another species,
who then rear their adopted young.
crown: The top of the head. [17]
CRP: See Conservation Reserve Program. [1035]
crus: The lower leg; in birds, supported
by the tibiotarsus bone. [114]
cuckoo: General term for birds in the large
family Cuculidae. The nearly 50 species
of Old World cuckoos are in subfamily
Cuculinae, and are all obligate brood
parasites. The familiar two-note chime
of the cuckoo-clock mimics the song
of one of these speciesthe Common
Cuckoo of Eurasia. In the New World,
cuckoos are in two subfamilies, Coccyzinae, whose members, all nonparasitic,
are termed New World Cuckoos and
include the Yellow-billed Cuckoo, Blackbilled Cuckoo, and Mangrove Cuckoo of
North America; and Neomorphinae (the
neomorphine cuckoos), whose members
are all Neotropical and include three
species that are obligate brood parasites.
Cuckoo-Roller: A crow-sized, arboreal
bird with a stout, broad bill; it is the only
member of its family (Leptosomatidae)
and is found only in Madagascar. [188]

Cornell Laboratory of Ornithology

cuckoo-shrikes: A diverse group of arboreal songbirds, many of which have a

shrike-like bill and are slender and (in
some cases) barred like cuckoos; however, they are related to neither group.
Together with minivets and trillers, they
make up the family Campephagidae,
found in the warmer parts of the Old
World. [196]
cup nest: A nest in the shape of a cup,
usually constructed of mud, or small
twigs and dried grass, with a depression
in the center to hold the eggs; built by the
majority of bird species. [830]
cursorial: Adapted for running. [E14]
cursorial theory (of the origin of avian
ight): A theory suggesting that the ancestors of birds evolved to y by rst running
along the ground, and then by jumping
and leaping, which was augmented by
the evolution of wings and feathers, which
eventually led to full ight. First proposed
by Samuel Williston in 1879. Also called
the Ground-Up Theory. [E14]
cytoplasm: The contents of a cell outside
the nucleus but within the cell membrane. [144]

dabbling: A foraging technique in which

a bird moves the beak rapidly on the surface of shallow water to pick up small
aquatic animals and plant materials; it is
used by dabbling ducks and a few other
species. [647]
dabbling ducks: Ducks (such as Mallards,
teal, wigeon, and pintails) that feed by
dabbling on the waters surface, in contrast to diving ducks, which dive under
water to search for plant material or
aquatic organisms. Dabbling ducks also
may tip bottom up and reach down under the water to obtain submerged food.
Also called puddle ducks. [647]
dark meat: See red bers. [57]
Darwins Finches: See Galapagos Finches. [160]
dawn chorus: The great amount of bird
song heard around dawn. At this time the
largest number of bird species are singing, and they sing more frequently, and
often more energetically and with more
variety than at other times of the day. Why
birds sing most at dawn, and in different
ways than they sing during the day, is not
known. [775]
dawn song: Bird song usually given only
during the early morning hours; it differs
from a species normal daytime song.
DDE: A stable, persistent, toxic organic
(1,1-dichloro-2,2-bis(pchlorophenyl)eth-ylene) formed in the

body by the metabolism of the organic
pesticide DDT (see separate entry). DDE
accumulates in fatty tissues and is excreted very slowly, and when concentrations become high it can cause death
or other toxic effects such as reproductive
failure resulting from eggshell thinning
(due to the disruption of calcium metabolism). Thin eggshells severely decreased
reproductive success in North American
raptors in the 1950s and 1960s, causing
populations of most raptor species to
plummet. [1052]
DDT: An organic pesticide (dichloro-diphenyl-trichloroethane) used commonly
in the United States from the mid-1940s
to the early 1970s to control Mexican
boll weevils, gypsy moths, mosquitoes,
and other insect pests. DDT is highly persistent in the environment and is taken in
by organisms and converted to DDE, a
toxic compound that accumulates in fatty
tissues and is excreted very slowly. DDT
was banned in the United States in 1972,
but is still used in other countries, including Argentina, Belize, Ecuador, Guyana,
Peru, and Mexico. See DDE for more information. [9125, 1052, 1053]
death rate: The number of individuals in
a species or population that die in a given
period of time; also called the mortality
rate. [949]
deciduous: Describing trees or shrubs
that lose their leaves during part of the
year. They generally drop their leaves
during the drier season (which in the
temperate zone is winter) to conserve
moisture, which might otherwise be lost
through evaporation. [9117]
decomposers: Level in a food chain or
web that consists of organisms (such as
earthworms, fungi, and bacteria) that
eat dead organisms and waste products,
breaking them down into basic nutrients
(such as oxygen, nitrogen, and phosphorus) and returning them to the soil for
plants to use. [9123]
decurved: Curved downward. Used to
describe beaks such as that of the White
Ibis. [117]
deferent ducts: Highly convoluted tubes
(also called vasa deferentia) that carry
sperm from the testes to (in birds) the cloaca; in birds, the lower portion of each
is enlarged to form a temporary storage
receptacle for sperm. [4128]
denitive plumages: Any of the plumages of a fully mature bird; they may change
seasonally, but do not change from year
to year as the bird ages. Some species,
such as gulls, large raptors, and pelagic
seabirds, take several years to reach their
denitive plumage. [330]
deective coloration: Conspicuous
markings found on otherwise cryptically
colored organisms; on birds, these mark-

ings show only in ight or when they are
ashed. They may have antipredator or
social functions. [364]
delayed plumage maturation: A situation
found in some species, in which one sex
remains in subadult plumage longer than
the other sex. [332]
dendrites: Rootlet-like extensions from
the cell body of a neuron; they usually
receive nerve impulses across a synapse
from other nerve cells, but may transmit
them as well. [432]
dens: The upwardly projecting knob or
peg of the axis (second cervical vertebra);
it ts into a hole in the ventral surface of
the atlas (rst cervical vertebra). [416]
density: See population density. [958]
density-dependent factor: A factor that
regulates population density in such a
way that the magnitude of its effect is
determined by the population density.
For example, population density is decreased by disease, which has a greater
and greater effect as population density
increases and thus rates of disease transmission go up. Other density-dependent
factors include predation, parasite levels,
and competition for resources. [969]
density-independent factor: A factor that
affects population density in such a way
that the magnitude of its effect does not
depend on the population density. Examples include severe weather, natural
disasters, and the failure of food supplies.
dentary: In birds, the bone forming each
side of the lower beak. [411]
deoxygenated blood: Blood whose red
blood cells carry very little oxygen. Also
called oxygen-poor blood, it is found in
all veins except the pulmonary veins and
venules. [481]
depolarizing material: A substance that
can be used to take a polarized light
beam and vibrate it in all directions, creating waves of all orientations, to form an
unpolarized beam. [594]
dermal papilla: The portion of a feather
papilla formed from dermal tissue; blood
vessels extend from the dermal papilla
into the developing feather, providing
nourishment. This core of tissue remains
in a feather follicle throughout a birds
life, ready to aid each round of feather development after a feather is lost. [326]
dermis: The inner layer of the skin; it lies
just beneath the epidermis and contains
blood vessels, muscles, and nerves.
descending aorta: The continuation
of the aorta (after it exits the heart and
curves) down through the body and toward the tail. [482]
determinate layers: Bird species that will

Handbook of Bird Biology


not lay additional (replacement) eggs if

one or more are removed from a clutch
during laying. Most determinate layers
will lay a new clutch if the entire clutch
is destroyed, however. For comparison,
see indeterminate layers. [890]
deterministic: In biology, describes
events whose occurrence and/or outcomes are inevitable, based on a certain
set of starting conditions. For comparison, see stochastic. [976]
dialect: A geographic cluster of similar
vocalizations (bird song, human speech,
or the sounds of other animals) that is
a consequence of those vocalizations
being learned. Dialects may exist over
very small or very large areas, depending
on the details of dispersal and learning.
diapsid: Describes a condition in which
the skull has two openings on each side
in the temporal region, posterior to the
eye socket. Diapsid skulls have this arrangement, and diapsid reptiles are those
reptiles that have diapsid skullsincluding thecodonts and their descendants
(including birds and crocodiles), snakes,
and lizards. Turtles are the only living
group of reptiles that are not diapsids.
differential exploitation: A situation in
which direct interactions among two
or more individuals that seek the same
resources are avoided because the individuals use the resources in slightly
different ways or use slightly different
resources. Differential exploitation may
result from the evolution of either morphological or behavioral differences
among individuals. [939]
dilution effect hypothesis: The idea that
colonial breeding may benet individuals because their large numbers can overwhelm the consumption capacity of local predators. [659]
dinosaur theory (of bird evolution): The
theory that birds evolved from theropods such as Compsognathus approximately 150 million years ago (proposed
by Thomas Huxley in 1868) or from
Dromaeosaurs such as Deinonychus
approximately 110 to 120 million years
ago (proposed by John Ostrom in 1973).
[E8, E10]
diopter adjustment ring: The ring on binoculars, usually on one of the eyepieces
but sometimes on the hinge post, that
allows the eyepieces to be focused independently to make up for the differences
in visual acuity between an individuals
two eyes. [230]
dip angle: The angle at which the magnetic eld lines around the earth contact
the earth. The dip angle is 0 degrees at
the magnetic equator, and approaches 90
degrees near the magnetic poles. [590]



Dir Due

direct benets hypothesis: One possible explanation for why females of

some species choose the males with
the most elaborate ornaments (such as
ornate plumage) to copulate with. The
explanation applies primarily to females
choosing nonpaternal sexual partners,
either (1) males for extrapair copulations,
or (2) mates in species in which males
do not provide parental care or other
resources for their offspring (such as territories with food or nesting sites). The
direct benets hypothesis suggests that
females may choose the most ornamented males because they are least likely to
infect the females with mites, a disease,
or some other afiction. Thus the female
gains reproductive advantages because
her health is not diminished by her mate
choice. [684]
direct tness: An individuals direct tness is the portion of its genes that is transferred into the next generation (and eventually beyond) through the production of
its own offspring. This contrasts with indirect tness, which is the portion transferred as a result of an individuals blood
relatives producing offspring. [686]
disjunct range: A range of a species or
population that is not continuous, but
rather is divided into geographically
separate areas. [950]
dispersal: In biology, usually the movement of individuals away from the area
where they were born, or away from
areas containing concentrations of individuals. [962]
displacement activities: Behaviors or actions that seem irrelevant or inappropriate to the current situation. For example,
a Herring Gull may stop to preen in the
middle of a territorial conict. Ethologists
hypothesize that displacement activities
occur because of conicting motivations
or indecision. Some displacement activities have become exaggerated and ritualized into displays. [632]
disruptive coloration: A type of cryptic
coloration with patches, streaks, or other
bold patterns of color that break up the
shape of the organism, catching the eye
and distracting the observer from recognizing the whole organism. [362]
distal: Away from the center of the body
(ngers are distal to the elbow) or from
the origin of the structure (the tip of a
feather is distal to its basewhere it is
attached). [14]
distraction displays: Displays in which
a bird or other animal feigns injury or in
some other way creates a highly noticeable fuss or disturbance, in order to shift a
potential predators attention away from
the birds nest or young. For example,
Killdeer give a broken-wing distraction
display by dragging and apping one

wing and uttering distress calls, while

slowly uttering along the ground away
from the nest or young. [653]
diurnal: Active during daylight. [931]
divergent evolution: A type of evolution
in which different populations of the same
species become increasingly distinct
from one another over many generations
(due to exposure to different ecological
factors), eventually diverging into two or
more new species. [157]
diving ducks: Ducks (such as Canvasbacks, Redheads, scaup, and goldeneyes)
that feed by diving under the waters surface to obtain aquatic plants or animals,
in contrast to dabbling ducks, which remain at the waters surface. [645]
DNA (deoxyribonucleic acid): The genetic material of all cellular organisms
and some viruses, forming the chromosomes of these organisms. A molecule
of DNA consists of two long strands of
nucleotides, held together by bonds
between nitrogenous bases on the two
strands, with the whole structure twisted
to form a double helix. The sequence of
the nucleotides is the genetic code,
which contains the instructions for making proteins, which in turn determine the
characteristics of an organism. [142]
DNA-DNA hybridization: A technique
used to determine the degree of similarity (in nucleotide sequence) between two
different samples of DNA. It is often used
to compare the DNA of two different species, to estimate how closely related they
are and to hypothesize their evolutionary
relationships. [146]
DNA ngerprinting: A technique by
which the nucleotide sequences of selected portions of the DNA of individuals of the same species are analyzed and
compared to determine how closely related the individuals are likely to be. Used
in criminology to determine if the DNA
of clues such as hair or semen match the
DNA of a suspected criminal; also used in
biological research to determine the relatedness among individuals whose family
histories are not knownsometimes to
avoid the breeding of closely related individuals of endangered species. [1076]
domed nest: A cup nest with a woven
overhead dome that likely helps to conceal the eggs or nestlings; built by meadowlarks, snipe, Ovenbirds, and others.
dominance hierarchy: A ranking system
of social status among members of a
group; often established and maintained
through displays and various aggressivesubmissive behaviors, including, on occasion, physical combat. Many dominance hierarchies are linearA dominates B, B dominates C, and C dominates
Dbut other arrangements exist. [624]

Cornell Laboratory of Ornithology

dorsal: Toward the back (the vertebral

column) of an organism. [14]
double-clutching: A technique in which
biologists remove one or several eggs
from the nests of indeterminate layers
and rear those eggs in the lab while the
birds lay replacement eggs and rear the
resulting young in the wild. Used by conservationists to increase the population
size in declining species such as California Condors and Peregrine Falcons.
down feathers: Soft, uffy feathers, typically lacking a rachis. Because the barbules lack hooks, the barbs do not cling
together, so they trap more air and thus
provide extra insulation. Some adult
birds have body downs under their contour feathers, and young birds have natal
down before molting into their juvenal
plumage. [316]
drafting: Driving closely behind another
vehicle and being pulled forward by air
currents moving back over the top of
the rst vehicle and swirling down and
forward, as well as by air currents swirling up and forward from below the rst
vehicle. Used by race car drivers or cars
following trucks on a highway; the energetic advantages are similar to those
experienced by a bird ying close to
and straight behind another bird, as in a
straight-line formation. [547]
drag: A force on a moving object, resulting from the friction between the object
and the uid (such as air) that it is moving
through. [516]
dromornithids: A group of giant, ightless birds that lived in Australia beginning
in the Tertiary period, but that became
extinct about 10,000 years ago during the
Ice Age. They are thought to be neognathous birds most closely related to the
Anseriformes (ducks, geese, and swans).
drumming: 1. Nonvocal sounds produced by woodpeckers banging on dead
trees or other resonant objects with their
beaks; most sound like short, emphatic
drum rolls, and are given by both sexes
to proclaim territory and attract mates.
[715] 2. A series of accelerating, nonvocal, mufed thumps at a very low
frequency, produced by male Ruffed
Grouse to proclaim territory and attract
mates. Produced by repeatedly bringing
the wings forward and upward so rapidly that with each stroke they compress
a parcel of air between the chest and
wings, creating a sound wave without the
wings and chest ever touching. Usually
performed while standing on a hollow
log, which acts as a resonating chamber
to amplify the sound. [715]
duet: Singing performance by two individuals. Avian duets usually are given by

a paired male and female who may sing
in synchrony, overlap one another, or
alternate their songs, depending on the
species. Most duetting species are tropical, and include some parrots, woodpeckers, antbirds, ycatchers, shrikes,
and wrens. [778]
dump laying: Laying an egg or eggs in
the nest of a conspecic (or sometimes,
a similar species). Females that dump lay
usually also build their own nests and
incubate their own eggs. Wood Ducks,
Hooded Mergansers, and some other
cavity-nesting waterfowl commonly
dump lay, sometimes resulting in large
numbers of eggs in one nest. Also called
dump nesting or egg dumping. [876,
duodenum: The U-shaped rst loop of
the small intestine, running from the
stomach to the jejunum. [4122]
dura: The outermost, toughest, and
most brous vascularized membrane
surrounding the brain and spinal cord.
Along with the other meninges, it provides sustenance and waste removal for
the cells of the brain and spinal cord,
which are not served by the circulatory
system. [436]
dust-bathing: Driving ne particles
through the feathers by rolling the body,
ufng the feathers, and wiping the head
and bill in a dusty area or by picking up
dust and throwing it over the body, after which dust is shaken or preened out;
important for feather maintenance and
removal of ectoparasites. [321]
dynamic pressure: The pressure of a
owing uid, or of movement through
a uid such as air or water. You feel dynamic pressure when wind blows against
your face. [512]
dynamic soaring: A type of ight in which
birds use the gradient in wind speed that
exists over the surface of the ocean to
travel for long distances without spending much of their own energy: the bird
glides down the gradient at an angle, then
turns and abruptly rises into the wind, using its momentum to gain height quickly,
then turns and glides down again, crossing the ocean in large zig-zags. Dynamic
soaring is used most by albatrosses and
other large pelagic birds with high-aspect-ratio wings. [544]

eardrum: See tympanic membrane.

eastern montane forest: North American
coniferous forest ecosystem dominated
by spruce and r trees and found below
alpine tundra but on the higher summits, ridges, and slopes of the Appala-

Dum Edg
chians from northern New England south
through Georgia and Alabama; it hosts
relatively few birds. [9116]
eclipse plumage: The set of dull-colored
feathers worn briey after the breeding
season by some adult birds, such as ducks.
In eclipse plumage, male ducks look like
females, which do not change much in
appearance. Eclipse plumage is acquired
by a complete molt after the breeding season, and is soon replaced through a partial
molt that produces the brighter colors of
the breeding plumage. [334]
ecological constraints hypotheses: A set
of hypotheses that each give an explanation for why certain individual birds
might forego their own breeding in a particular breeding season, instead acting
as helpers at the nest of other breeding
pairs (usually their parents or other close
relatives). The ecological constraints hypotheses focus on the possible costs to
young birds of dispersing from their natal
territory; examples of these hypotheses
include (1) few vacant territories of good
quality may be available, (2) few suitable
breeding partners may be available, and
(3) the birds may have little chance of
reproducing successfully until they gain
parenting experience. [689]
ecological isolating mechanisms: Structural, physiological, or behavioral adaptations that have evolved in species that
have very similar niches, and that allow
the species to divide up the resources in
various ways (such as using slightly different resources or foraging in different
parts of the habitat), and thus to coexist.
For example, certain wood-warbler species can coexist in spruce forests of northeastern North America even though they
all eat similar small insects, because they
forage in different parts of trees. [9103]
ecological niche: See niche. [9102]
ecological release: An expansion of the
niche of certain populations of a species,
such that a greater breadth of resources
such as habitat and food are used, in areas where interspecic competition is
lower, as on islands. (In these situations,
the niche is released.) [9104]
ecological succession: The process by
which one association of plants and animals is replaced by another, then that one
is replaced by another, and so on until
the climax community for that area is
reached. The types of communities, and
the order in which each succeeds the previous one, is fairly predictable for a given
habitat and geographical region. Primary
succession begins on a substrate, such as
rock, sand, or lava, that has never before
supported a community. The process by
which a lake gradually lls in to form
a bog community also is considered a
form of primary succession. Secondary

Handbook of Bird Biology


succession begins with bare soil or an

existing community. [9109, 9110]
ecology: The study of the relationships
between organisms and their environment. [91]
ecoregional planning: Careful, biologically driven planning at the level
of ecoregions, with the ultimate goal of
preserving the species and ecosystem
processes (for example, re and pollination) that occur within each ecoregion.
Ecoregional planning involves geographically delineating ecoregions; cataloging
the ecosystems within each ecoregion;
identifying and mapping the most important species, communities, and habitats;
determining potential threats to species
and sites; and then prioritizing species,
communities, and the key habitats as to
which require the most urgent conservation action. Ecoregional planners then
set conservation goals, outline plans, and
monitor the results. [1086]
ecoregions: Regions of the world that are
biologically distinct in terms of climate
conditions, topography, soil types, and
plant communities; also called physiographic regions. [1086, 1087]
ecosystem: Both the living and nonliving
components of a particular area (including the physical surroundings), as well as
the ecological processes that bind them
all together (such as decomposition, soil
erosion, and water and nutrient cycling).
Ecosystems may be small (for example,
the ecosystem of a rotting log) or large (a
deciduous forest ecosystem). [9109]
ecosystem management: Understanding and maintaining entire ecosystems
(see separate entry), instead of focusing
on particular species or habitat types.
ecotone: A zone of transition between
two ecosystems, such as the oak savanna
ecotone between the eastern deciduous
forests and prairie grasslands of North
America. Ecotones host a greater number of species than either adjacent ecosystem, because some species in each of
the two adjacent ecosystems frequent the
transition zone, and because some species prefer the greater variety of resources
found at habitat edges and thus live specically in those areas. [993]
ectoparasites: Parasites, such as ies,
ticks, eas, lice, and mites, inhabiting the
exterior of a hosts body. [319, 323]
ectothermic: Describes organisms that
must rely on sources of heat outside their
own bodies to keep warm; also called
cold-blooded. [11]
edge effect: The tendency for areas near
the edge of a habitat patch to differ from
areas near the center in a number of different ways. For example, areas where
two habitats meet fairly abruptly (such as




a forest/eld boundary) often host a higher number of species than do either of

the two adjacent habitats. Other ways in
which the edge of a habitat patch may differ from the center include changes in microclimate, such as amount of sunlight or
humidity; increased habitat disturbances,
such as re and wind damage; and higher
numbers of introduced plants or animals
(because these species generally invade
from the edge). Forest birds breeding near
edges may experience higher rates of predation and/or brood parasitism than birds
breeding near the center, because predators such as raccoons, squirrels, and Blue
Jays and parasites such as Brown-headed
Cowbirds are more common near edges.
[993, 1073]
egg: 1. The ovum; the female reproductive
cell sometimes called the egg cell, both
before and just after it is fertilized by a
sperm cell. [4130] 2. The hard-shelled
structure laid by birds, containing the
embryo, yolk, and white. [4130]
egg tooth: A short, pointed, calcareous
structure on the tip of the upper beak (and
sometimes the lower beak as well) that
develops in bird embryos shortly before
hatching; the embryo rubs and pounds
the egg tooth against the inner wall of
the eggshell to break it open and hatch.
The egg tooth sloughs off or is resorbed
by the growing chick within a few days
after hatching. [8104]
electrophoresis: A method of separating
large, charged molecules of different
lengths or charges (DNA fragments or
proteins treated to carry a charge) by their
rate of movement through a thin slab of
gel in an electric eld. [143]
elephantbirds: A family (Aepyornithidae)
of huge, ightless ratites that lived on the
island of Madagascar beginning 10 to 20
million years ago, but were exterminated
about 2,000 years ago by human activity.
The tallest stood about 10 feet (3 m) and
weighed about half a ton. [E23]
elliptical wings: Short, broad wings having a low aspect ratio; they allow great
maneuverability, but do not promote
efcient or rapid ight. Elliptical wings
are common in birds that live in forests,
woodlands, or shrubby areas, such as
crows, grouse, quail, and most songbirds. [537]
emarginate tail: Tail shape in which the
rectrices become slightly longer from the
inside out. Also called notched. [119]
embryo: A developing young animal that
is still inside its egg or mother; in some
animals, especially mammals, refers
only to the earlier developmental stages.
emigration: The movement of individuals
out of a population. [949]
Emu: A large, ightless ratite of Australia;

it is the only member of its family (Dromiceidae) and is the second largest living
bird, next to the Ostrich. [193]
enantiornithine birds: See opposite
birds. [E20]
Endangered Species Act (ESA): Federal
law passed in the United States in 1973
that commits the government to take action to prevent the extinction of native
species and to protect their habitat. It
also establishes a procedure to develop
a list of threatened and endangered species, identify their critical habitat, and
develop and carry out Recovery Plans.
[1023, 1091]
endemic: Found only in a particular region; describes a species or other taxonomic group. For example, kiwis are
endemic to New Zealand. [170]
endocrine glands: Structures that secrete
hormones directly into the blood; the
hormones are carried to other parts of the
body, where they stimulate or regulate
the activities of other glands or organs.
The major endocrine glands of birds are
the pituitary, thyroids, parathyroids, ultimobranchials, adrenals, gonads, pancreas, pineal, thymus, and cloacal bursa.
endocrine system: Organ system that
acts with the nervous system to initiate,
coordinate, and regulate body functions, including reproduction and development. It consists of the endocrine
glands and their secretions, called hormones. [469]
endolymph: The uid that lls the structures forming the membranous inner labyrinth of the inner ear: the cochlear duct,
the semicircular ducts, and the utriculus
and sacculus. [457]
endothermic: Having the ability to
generate ones own body heat through
metabolic processes; also called warmblooded. Only birds and mammals are
endothermic. [11, 913]
entoglossal: The bone (part of the hyoid
apparatus) that supports the tongue; also
called the tongue bone. [414]
environment: The surroundings of an
organism, including physical features,
chemical and energetic factors, and other living organisms. [97]
environmental stochasticity: The tendency of nearly all environments to experience many random (or at least unpredictable) events, such as natural disasters
or severe weather. If a population is small
and unlucky enough to be struck by
one of these events, it may be entirely
wiped out. [977]
enzyme: A protein that catalyzes (assists)
a biochemical reaction without being
consumed in the reaction. [141]
epaulettes: Patches of colored feathers on

Cornell Laboratory of Ornithology

a birds shoulders, such as the red feathers

on a Red-winged Blackbird. [35, 673]
EPC: See extrapair copulations. [679]
epidermal collar: Structure formed during feather development as a feather papilla elongates: the outer cells harden,
fuse, and form a ring or collar of epidermal tissue surrounding the dermal
portion of the original papilla. The collar
cells multiply to produce most structures
in the developing feather. [326]
epidermis: The outer layer of the skin; it
protects the inner layerthe dermis
and does not contain blood vessels. In
birds, it gives rise to the feathers and to
the horny sheath covering the bill, legs,
and feet, including the claws. [326]
epiphytes: Plants that grow on other
plants but, in contrast to parasitic plants,
use their roots for attachment rather than
to obtain nutrients from their support
plant; examples include many bromeliads (air plants) and orchids, and a few
cacti. [988]
epizootic: 1. A disease that spreads
quickly among crowded animals, such as
viral enteritis (duck plague), which may
kill hundreds or thousands of waterfowl.
[969] 2. Term used to describe a disease that spreads quickly among animals
when they are highly crowded. [969]
epoch: A unit of geological time. Successive epochs make up a period. [1113]
equilibrium theory: Theory proposing
that the number of species on an island
at any one time represents a balance between the number of new species colonizing the island (immigration) and the
number of species becoming extinct. A
consequence of this relationship is that
smaller islands tend to have fewer species because (1) they tend to have lower
immigration rates (they are less likely to
be discovered by colonists), (2) they
have higher extinction rates (the populations are smaller and thus more likely to
go extinct due to stochastic factors, and
the fewer resources and reduced habitat
diversity are more likely to lead to competitive exclusion), and (3) they typically
contain a lower diversity of habitat types
than larger islands, thereby providing
niches for fewer kinds of species. This
theory applies only in situations in which
speciation is not a major source of new
species, for example over short periods
of time, or on islands too small or too far
from other islands to permit speciation
through geographic isolation. [995]
era: A unit of geological time. Eras are
divided into periods. [1113]
erythrocytes: See red blood cells.
ESA: See Endangered Species Act.
[1023, 1091]

esophagus: Thin, straight muscular tube
carrying food from the pharynx to the
stomach. It is lined with mucus-secreting
glands that moisten food to ease its passage, but contains no digestive glands.
ethology: The study of animal behavior,
primarily from a proximate approach,
by comparing similar behaviors in related species to understand how certain
behaviors evolved, and by investigating
releasers and instinctive behaviors and
the underlying physiological processes.
Some of the earliest animal behaviorists
(primarily Europeans) were concerned
mainly with ethology, and the two pioneers of the eld, Konrad Lorenz and
Niko Tinbergen, laid the groundwork
for animal behavior studies carried out
today. [67]
eustachian tube: See auditory tube.
eutrophication: The changes in a lake,
estuary, or slow-moving stream when
it receives excess plant nutrients, especially nitrates and phosphates. Some nutrient input occurs naturally, through the
erosion of soil and run-off from adjacent
land. In many cases, however, human
activity greatly increases the rate of nutrient input or adds new sources, such as
discharges from industries and sewage
treatment plants; when human activity
is involved, the changes are sometimes
termed cultural eutrophication. One important change is the dramatic increase
in the growth of plants and (especially)
algae, which can choke out native plants.
Then, as the masses of plants and algae
die and sink to the bottom they decompose, depleting the water of oxygen, and
thus few aquatic organisms can continue
to survive. In addition, some algae may
produce toxins, dyes, or odors that decrease water quality. [1056]
eventual variety: Pattern of singing in which
a bird repeats one song type many times before switching to a different type, which it
then repeats many times, and then switches
to another type, and so on. [786]
evolution: A change over time. The evolution of living things is the set of cumulative changes in the characteristics of a
species or population over successive
generations that result from natural selection acting on the genetic variation
among individuals. [134]
excavators: In avian biology, bird species
that dig their own nest cavities or tunnels, either in sandy soil (such as Belted
Kingshers, Bank Swallows, bee-eaters,
mot-mots, and Crab Plovers) or in wood
(woodpeckers). [839]
excretory system: Organ system consisting of the kidneys and their ducts, the
ureters (in the rheas of South America,

Eso Fea
but in no other birds, a urinary bladder
is also present); it removes toxic nitrogenous wastes from the blood by producing, storing, and excreting urine. In birds,
the urine is composed of uric acid. Also
called the urinary system; it is often considered together with the reproductive
system as the urogenital system. [4125]
exit pupil: The space through which
the light beam exiting the eyepiece of
binoculars or a telescope passes; this
is, in effect, the hole through which the
observer looks. The diameter of the exit
pupil is calculated by dividing the size of
the objective lens by the magnication;
the larger the exit pupil, the brighter the
image. [235]
extensor: A muscle that pulls one bone
away from another bone. [427]
external ear: The portion of the ear containing the external ear canal and the eardrum (tympanic membrane. [456]
external ear canal: The channel through
which sounds enter the ear; it leads from
outside the body to the eardrum. [456]
extinction events: Large-scale extinctions of many species; they have occurred periodically throughout Earths
history. [1113]
extra-embryonic membranes: Membranes that protect and nourish the growing embryo, but do not become part of
the adult body. The four main ones in the
avian egg are the yolk sac, amnion, allantois, and chorion. [865]
extrapair copulations (EPCs): Copulations with birds other than ones mate.
eyeball: The eye; in birds it is at to tubular, in contrast to the spherical eyeball of
mammals. Its outer layer is the sclera, and
the entire eyeball sits in the socket and is
protected on its exposed side by the eyelids and nictitating membrane. [447]
eyebrow stripe: A distinctively colored
line running from the upper beak toward
the back of the head, located ventral to
the boundary of the forehead and crown;
also called the superciliary line. [17,
eyeline: A distinctively colored line that
passes through the eye. [210]
eye ring: A circle of distinctively colored
feathers or skin surrounding the eye.

facial disc: Flat, relatively round, forwardfacing part of the head of owls; it probably
funnels sounds into the birds ear openings. Also spelled facial disk. [455]
facial nerve: The seventh cranial nerve;
it carries motor signals to muscles that

Handbook of Bird Biology


protrude the tongue, lower the lower

beak, constrict the neck, and (in birds)
tense the columella ear bone. It also may
carry some taste sensory input from the
tongue. [441]
facultative partial migration: See partial
migration. [556]
fairy-bluebirds: Two species of arboreal
songbirds endemic to the Oriental zoogeographic region; they are named for
the brilliant blue and black plumage of
the males, and feed primarily on gs and
other fruits. Together with leafbirds, they
make up the family Irenidae. [189]
fairywrens: An Australasian family (Meluridae, 26 species) of cooperatively
breeding, wrenlike birds with long,
cocked tails. [195]
false sunbirds: Members (with asities)
of the family Philepittidae, endemic to
Madagascar. These two species of suboscine passerines feed on insects and
nectar. [188]
family: Level of classication of organisms above genus and below order;
similar genera are placed in the same
family. The scientic names of bird families end in idae (for example, Corvidae). [152]
FAP: See xed action pattern. [68]
fascia: Connective tissue binding together hundreds or thousands of muscle
bers to form a skeletal muscle; it may be
in the form of bandlike tendons or broad,
shiny sheets called aponeuroses. [427]
fat bodies: In birds, yellowish fat deposits laid down just under the skin, usually
in individuals storing fat in preparation
for migration; the most conspicuous fat
bodies lie over the abdomen and in the
depression formed anterior to the breast
muscles where the clavicles fuse to form
the wishbone, and are visible in a handheld bird with the feathers parted. Researchers often use the degree of fat accumulation in a migrant as an indication
of its energetic condition and potential to
continue migration. [564]
feather comb: See pectinate claw.
feather papillae: Small bumps covering
the surface of the skin of birds during
embryonic development. They consist
of a core of dermis and a covering of epidermis, and each will eventually form an
embryonic feather. Also simply called
papillae. [326]
feather sheath: A thin, cylindrical tube
of keratin surrounding and protecting a
developing feather. It eventually breaks
open to let the mature feather unfurl.
feather tracts: Areas of a birds skin
where feathers are attached; also called
pterylae. [32]



Fec Fur

fecal sac: A tough, exible bag enclosing

the feces of most passerine nestlings; it
allows the parents to remove and dispose
of the feces more easilyparents sometimes grab the fecal sacs as they emerge
from a nestlings cloaca. Many parents
carry the sacs some distance from the
nest and drop them, but others consume
them. [8136]
fecundity: See birth rate. [949]
female-defense polygyny: Mating system
in which males compete ercely for control of clusters of nesting females. In some
species, such as Montezuma Oropendolas of Central America, a male dominance
hierarchy results with the top few males
securing most of the matings. Because
males do not help to rear the young, this
system may evolve when male parental
care is less important to the survival of the
young than are safe nesting sites or rich
food supplies. [674]
femur: Bone that supports the upper hind
limb (thigh) of many vertebrates, including birds and humans. [114]
bula: The thinner of the two lower hind
limb bones in many vertebrates, including humans; in birds, the bula is reduced
and present only as a thin, needlelike
bone running two-thirds of the way down
the side of the tibiotarsus. [114, 425]
eld of view: 1. The width of the area
visible (usually at 1,000 yards from the
observer) through binoculars or a telescope. If the eld of view is labeled in
degrees, multiply degrees by 52.5 to get
the width in feet. 2. The view attained
by a particular species, due to the placement of its eyes; also called an organisms
visual eld. [237, 451]
loplumes: Hairlike but relatively stiff
feathers having a rachis but few or no
barbs (any barbs are present only at the
tip). In the skin next to their follicles they
have sensory receptors that allow them
to monitor movement within the feather
coat. [317]
tness: An individuals degree of success
at contributing its genes to the next generation; often measured as the number
of its offspring that survive to reproduce.
[135, 97]
xed action pattern (FAP): A behavior
that occurs in complete form each time
the animal encounters the releasing
stimuluseven upon the animals rst
exposure to that stimulus. An FAP may
be a simple or complex behavior, or a
series of behaviors, but once begun, it is
played out to the end regardless of any
response that occurs or any intervening
stimuli. [68]
anges: See oral anges. [343, 8107]
ank: The side of a bird, dorsal and caudal to the leg. [16]

edging: Term commonly used to describe the time at which nestlings that
are reared in the nest leave the nest, even
though their ight abilities may not yet
be well developed. But, the term is sometimes used to describe the time at which a
young bird has nished acquiring its rst
complete set of ight feathersgenerally
the time at which it is capable of ight.
The term is used less often in precocial
species that leave the nest shortly after
hatching, but sometimes it refers to the
time at which they begin to y. Fledging may also be used to refer to the process of reaching the moment of edging.
edging period: The period of time from
hatching to the moment of edging (see
separate entry). [8116]
edgling: A young bird that has recently
edged (see separate entry for edging).
exor: A muscle that pulls one bone toward another bone. [427]
ight feathers: The remiges of the wings
and rectrices of the tail. [111, 112,
ight songs: Songs given by birds during ight; they are particularly common
among birds of open areas, such as grasslands and the tundra, where few perches
are available. Singing from higher up generally increases the distance over which
a song can be heard. Nevertheless, some
species that do have ample perches, such
as the forest-dwelling Ovenbird, also give
ight songs; these usually begin with a
jumble of notes that appear to draw attention to the singer, and then proceed with
the birds normal song. Their function is
unknown. [784]
oaters: Animals, generally males, that
do not hold territories or form pair bonds,
but cruise around areas containing territorial individuals, waiting for a chance to
take over a territory or sneak a copulation
with a paired bird. [970]
owerpeckers: A family (Dicaeidae, 43
species) of small, busy, noisy songbirds,
primarily of the Oriental zoogeographic
region, that forage high in trees on berries, nectar, and insects. [190]
follicle: A small, epidermis-lined pit in
the skin of a bird, from which a feather
emerges and to which it is attached. [32,
food chain: The sequence in which organisms in an ecosystem feed upon other
organisms. Most food chains consist of
producers, consumers, and decomposers. A food chain is a fairly linear and
simplistic model of feeding relationships
in communities, which are more realistically represented as food webs because
of the numerous interconnections among
species and levels. [9123]

Cornell Laboratory of Ornithology

food web: See food chain. [9123]

foot: In birds, refers to the portion of the
leg distal to the tibiotarsus bone, and
has two sections. The upper section is
supported by the tarsometatarsus bone,
which does not touch the ground when
the bird walks; the lower section consists
of the phalanges of the toes, upon which
the bird walks. [114]
foramen triosseum: An opening at a birds
shoulder joint, formed by the junction
of the scapula, coracoid, and clavicle
bones. This hole acts as part of a pulley
system that allows the force of the supracoracoideus muscle to be redirected:
because the tendon of the supracoracoideus passes through this hole, the muscle
can be located ventrally yet still raise the
wing. Also called the triosseal canal or
supracoracoid foramen. [420, 56]
forebrain: The anterior portion of the
brain; it consists of the two cerebral
hemispheres with the olfactory lobes at
their anterior ends. [437]
forehead: The front of the head, from the
crown to the base of the bill. [17]
forked tail: Tail shape in which the rectrices become abruptly longer from the
inside out. [119]
formation: An ordered arrangement of a
group of birds in ight, such as V-shaped
ocks of geese or single lines of Brown
Pelicans and cormorants. [547]
frequency: Also known as pitch, the
frequency is the rate at which a sound
causes the air through which it is moving
to compress and thin (one compressing
and thinning is called one cycle), and is
measured in cycles per second, or Hertz
(Hz). The more cycles per second, the
higher the frequency, and the higher the
pitch. [74]
fright molt: See shock molt. [338]
frogmouths: A family (Podargidae, 14 species) of nocturnal forest birds of the Oriental and Australasian regions that resemble
their smaller nightjar relatives in both cryptic appearance and behavior. [191]
frontal plane: A plane, usually horizontal,
through an organism, dividing the body
into dorsal and ventral portions. [14]
frugivorous: Feeding mainly or exclusively on fruits. [181]
functional response: A change in the
amount of a certain type of prey taken
by a predator, as a result of a change in
the population density of that prey. For
example, as insect populations in an
area increase during spring, American
Kestrels may begin to eat more insects
and fewer voles. [925]
furcula: V-shaped bone of the pectoral
girdle of birds, formed by the fusion of the
right and left clavicles with a small inter-

clavicle bone. The furcula is also called
the wishbone. [419]
fusiform: Describes an egg that is slightly
longer than subelliptical; also called biconical or long subelliptical. [873]

Galapagos Finches: A group of 15 nch

species in the family Emberizidae living
in and near the Galapagos Islands. They
are a classic example of adaptive radiation, as they have a wide array of beak
sizes and shapes and are all thought to
have evolved from a common ancestor.
Also called Darwins Finches. [160]
gall bladder: Small organ for storing bile;
it is located under the liver and is not
present in all birds. [4124]
gallinaceous birds: Grouse, quails, turkeys, pheasants, and all other birds in
the order Galliformes; includes domestic
chickens. [166]
gametes: The ova (egg cells) and sperm
cells. [4134]
ganglia (singular, ganglion): Aggregations
of nerve cell bodies; they form nerve centers outside the central nervous system
(in the peripheral nervous system). [436,
gaping: In avian biology, begging behavior of young birds that begins shortly after
hatching in which they open the mouth
widely; may be accompanied by a begging call. Given by altricial young and
those precocial young whose parents
feed them. [8107]
gas exchange: The movement of gases between an organism and the environment;
for example, in the lungs of many organisms including birds, the blood takes up
oxygen from the air and discharges carbon dioxide and water. [489]
gastric cuticle: Leathery or sandpaper-like
material that forms the lining of the gizzard; it is a combination of carbohydrate
and protein secreted by glands in the wall
of the gizzard. Also called koilin. [4119]
gastrointestinal tract: See alimentary
canal. [4103]
gene: The sequence of base pairs within
a molecule of DNA that codes for one
specic protein. [141, 4134]
gene ow: The movement of genetic material between populations. In mobile
animals, gene ow generally occurs as
individuals emigrate, immigrate, or breed
with individuals from other populations. In
organisms such as plants and fungi, gene
ow occurs as spores, pollen, or seeds are
carried by water, wind, or animals. [153]
gene pool: All the genes existing in a
population at a given time. [622]
generalist: In biology, an organism that

Fus Goo
is able to use a wide range of some type
of resource; for example, animals with
generalist diets eat many different types
of foods. [95]
genetic bottleneck: The loss of genetic diversity experienced by most populations
as they become very small. Such loss occurs for three main reasons: (1) some alleles are lost simply by chance, when the
only individuals that possess them die, (2)
the rate at which new alleles arise in the
population declines, because there are
fewer individuals in which new mutations
can occur, and (3) inbreeding occurs more
frequently because fewer genetically dissimilar individuals are available as potential reproductive partners. [1075]
genetic structure: The relative proportions of individuals with different genetic
typesusually noted for a given population. [980]
genital system: The reproductive system.
genus (plural, genera): Level of classication of organisms above species and
below family. Genus is always capitalized, and is underlined or printed in
italics. [147]
geographic range: The geographic area
within which a species or population
generally remains at a particular time of
year; a species may have different breeding and nonbreeding ranges. Also called
the range. [219, 949]
germinal spot: In avian biology, the lightcolored site on the egg yolk where the
embryo will eventually develop. The germinal spot sits atop a cylinder of lightcolored yolk that stretches from the yolks
core to its surface. [863]
gizzard: The lower part of the birds twopart stomach; it is rounded and has a
tough lining and thick, muscular walls,
often with internal ridges. It grinds and
softens foods, and in birds that eat seeds,
the gizzard has more muscular walls than
in birds that eat meat. Seed-eating birds
may eat grit or small stones, which reside in the gizzard to aid in grinding. In
birds that eat sh or other meat, the gizzard molds indigestible material, such as
bones and feathers, into compact balls
(pellets) that are then ejected through the
mouth. [4118]
gleaning: A foraging technique in which
a bird takes insects and other small invertebrates from the surface of vegetation
or other substrates. In perch gleaning,
practiced by many wood-warblers and
other species, a bird grabs prey without
ying from its perch. In sally gleaning,
practiced by birds such as Red-eyed
Vireos, chickadees, titmice, and some
small ycatchers, the bird sits still and
watches the surrounding vegetation until
it sees an insect move, then ies out and

Handbook of Bird Biology


grabs it from the surface. In hover gleaning, practiced by kinglets, phoebes, and
Great Crested Flycatchers, among others,
the bird hovers while taking food from the
surface of vegetation. [644]
glenoid fossa: In birds, a cup-shaped
depression formed where the coracoid
and scapula meet; it receives the rounded
end of the humerus, forming a ball-andsocket joint that enables the humerus to
rotate freely around the shoulder joint.
glial cell: See neuroglia. [436]
gliding: Unpowered ight (no thrust is
provided) in which the ying object loses
altitude. In a bird or other animal, ying
without apping the wings or limbs, while
losing altitude (as compared to soaring,
in which the animal rises). [536]
globular nest: A spherical dome nest
with a top that completely encloses the
nest; usually entered through a hole on
the side. Examples include the nests of
Cactus Wrens, Black-billed Magpies,
and Southern Penduline-Tits. [836]
glossopharyngeal nerve: The ninth cranial nerve; it carries sensory and motor
information between the brain and the
tongue, pharynx, esophagus, and throat.
It also carries motor output to the salivary
glands. [442]
glottis: Small, slit-like opening to the
larynx; it is surrounded by eshy folds
whose muscles regulate the passage of air
into the respiratory system. [491]
glycogen body: A gelatinous mass of
neuroglial cells rich in the nutritive sugar
glycogen; it is located in the rhomboid
sinus, and its function is unknown. The
rhomboid sinus and glycogen body are
unique to birds. [439]
gonads: The primary sexual organs, the
testes and ovary, which produce, respectively, sperm and eggs. They also are
endocrine glands, secreting the sex hormones testosterone, estrogen, and, from
the ovary only, proges-terone. [475]
Gondwanaland: The southern land mass
formed 200 million years ago when Pangea split into two large land masses. It
consists of present-day South America,
Africa, Madagascar, India, Australia,
New Zealand, and Antarctica. [168]
good genes hypothesis: One possible explanation for why females of some species
choose the males with the most elaborate
ornaments (such as ornate plumage) to
copulate with. The explanation applies
primarily to females choosing nonpaternal sexual partners, either (1) males
for extrapair copulations, or (2) mates in
species in which males do not provide
parental care or other resources for their
offspring (such as territories with food or
nesting sites). The good genes hypothesis



Gra Haw

suggests that the most ornamented males

have genes that increase their own survival in some way (for example, they may
have greater skill at foraging, avoiding
predators, or obtaining good territories),
and that females choose them because
then their own male and female offspring
may inherit those traits and have an increased chance of surviving and producing offspring of their own. [684]
graduated tail: Tail shape in which the
rectrices become abruptly longer from
the outside in. [119]
granivorous: Seed-eating. [927]
gravity: The attractive force between two
masses of matter; this force, for example,
tends to draw objects toward the center
of the earth. [510]
gray matter:
Darker-colored tissue
(compared to white matter) that makes
up much of the brain and spinal cord. It
consists of numerous nuclei, which are
collections of nerve cell bodies, and is
found at the core of the spinal cord and in
the outer areas of the brain. [439]
grazing: A foraging technique in which
an animal bites off clumps of grass or other vegetation; used by geese, antelope,
and others. [646]
greater coverts: The feathers partly overlying each remex on the upper surface of
the wing. A greater primary covert overlies each primary feather, and a greater
secondary covert overlies each secondary feather. [112, 113]
gross primary productivity: See primary
productivity. [987]
ground-rollers: A family (Brachypteraciidae, 5 species) of solitary, terrestrial insect-eaters with stout bills, short wings,
and moderately long legs and tails. They
are endemic to Madagascar and are declining in number dramatically. [188]
ground speed: The speed of a moving object (such as a ying bird) in relation to
the ground; it is equal to the birds speed
with respect to the air (the air speed) plus
or minus the wind speed, depending on
the direction the bird is ying with respect to the wind. [568]
ground-up theory (of the origin of avian
ight): See cursorial theory. [E14]
group selection: Theory proposed by V.
C. Wynne-Edwards in 1962 suggesting
that different groups of individuals might
experience differential survival based on
variations among the groups, and that this
might be one way in which natural selection workedpassing down through the
generations a greater proportion of the
genes of the groups with higher survival
rates. Theorists have basically disproved
the idea as originally presented, but are
still exploring whether some other form
of group selection might be plausible.

guano: The accumulation of seabird
droppings, particularly at a breeding
colony, dried to a hard, crusty state.
Guano is often mined and used as fertilizer. [1104]
guild: In avian biology, a group of ecologically similar (but not necessarily related) birds that use the same resources
in similar ways. For example, Henslows
Sparrows, Grasshopper Sparrows, Eastern Meadowlarks, and Bobolinks all nest
on the ground in extensive grasslands,
and thus are considered to be in the same
grassland nesting guild. [9102]
guineafowl: Gregarious, chicken-like
birds with distinctively spotted and
striped plumage; they are often domesticated or found in zoos. The six species are
endemic to Africa and form the subfamily Numidinae of the family Phasianidae.
gular uttering: Opening the bill wide
and vibrating the thin, expansive gular
membranes of the throat, in order to dissipate heat. This cooling method is used
by pelicans, cormorants, herons, owls,
and nighthawks. [4153]
gular region: Upper part of the throat, just
below the chin. [17, 18]
gut: See alimentary canal. [4103]

habitat: The physical surroundings in

which an organism lives. It consists of
physical factors, such as light, temperature, and moisture, as well as living organisms, such as plants and animals. Habitats
are often characterized by a dominant
plant type or physical feature, such as a
grassland habitat or stream habitat. [98]
habitat conservation plan (HCP): A plan
that must be submitted to the U. S. Fish
and Wildlife Service by anyone who applies for a permit to destroy endangered
species or their habitats (as allowed under
a 1982 amendment to the Endangered
Species Act). The plan must specify the
steps that the applicant will take to minimize the number of individuals killed
and to minimize the impact on the species as a whole, and also must explain
why other alternatives are not feasible.
habitat fragmentation: The process by
which a large, continuous habitat is
broken into a number of small, isolated patches by activities such as development, logging, or farming. [997]
habitat generalists: Species that can live
and breed successfully in a wide range
of different habitats. If all else is equal,
these species are less likely to go extinct
than species that can live and breed only

Cornell Laboratory of Ornithology

in one or a few specic types of habitats

(habitat specialists). [1059]
habitat imprinting: The process by which
a young animal, especially a bird, learns
the characteristics of appropriate habitat by observing its surroundings while it
is still living within its parents territory.
When adult birds eventually settle on
their own territories, most choose to live
and/or breed in areas similar to those in
which they were raised. [911]
habitat richness theory: The idea that
smaller islands generally hold fewer species than larger islands because they tend
to have fewer different habitats, and thus
species immigrating to smaller islands are
less likely to nd suitable habitats than
those colonizing larger islands, and are
more likely to perish (or leave). [996]
habitat specialists: Species that live and
breed successfully in only one or a few
specic types of habitats. If all else is
equal, these species are more likely to
go extinct than species that can live and
breed successfully in a wide range of
habitats (habitat generalists). [1059]
habitat specicity: How wide a range of
different habitat types in which a given
species can live and breed successfully.
habituation: The permanent loss of a response as a result of repeated stimulation
without reward or punishment; learning
to ignore unimportant stimuli. [69]
hacking: The technique of introducing
young, captive-raised birds of prey, especially falcons, to appropriate habitat
by releasing them from an enclosure that
serves as an articial nest and in which
biologists continue to place food until
the bird has learned to hunt on its own.
hallux: The rst toe, composed of two
phalanges. In nearly all birds, it points
backward. The hallux is well developed
in perching birds and is reduced or absent in many running birds. [425, 426]
Hamerkop: A stork-like water bird whose
shaggy, crested nape and stout, tapering
bill make the head appear hammershaped. It builds an enormous mound
nest. It is endemic to Africa, and is the
only member of its family, Scopidae.
hand: See manus. [19]
hatching: Emerging from the egg. A
clutch may hatch synchronously (all at
about the same timesee synchronous
hatching) or asynchronously (over a period of several dayssee asynchronous
hatching). [688]
hatchling: A newly hatched animal.
Hawaiian Honeycreepers: A group of 32
species, many of them extinct, in the sub-

family Drepanidinae, within the family
Fringillidae. These small, colorful birds
have a wide array of beak shapes and are
a dramatic example of adaptive radiation,
as they all are thought to have evolved
from a common ancestor. [162]
hawking: A foraging technique in which
a bird sits very still on a high or exposed
perch, and when it sees an insect, ies
out and snatches it in midair, returning
to the same or a nearby perch; used by
many ycatchers, kingbirds, bee-eaters,
waxwings (sometimes), and some woodpeckers, among others. [644]
HCP: See habitat conservation plan.
heading: The compass direction in which
a bird is pointing its beak and propelling
itself through the air. Because of crosswinds, the heading may not be the actual
direction that the bird is progressing with
respect to the ground. Also applies to other ying animals and aircraft. [568]
heel pad: Calloused enlargement of the
upper end of the tarsus (at the heel), found
in the nestlings of many cavity nesters,
such as woodpeckers and trogons. It is
thought to reduce abrasion of the tarsus
from the rough lining of the nest cavity.
helpers at the nest: Adult birds that are
not currently breeding themselves, but assist other breeding pairs (usually, but not
always, their relativesespecially their
parents) in rearing their offspring. Some
helpers are birds whose own breeding
attempts have failed, whereas others are
unpaired birds or those without territories; helpers often breed on their own in
subsequent years. Bird species in which
helpers are common are called cooperative breeders. [688]
hepatic portal system: Pattern of circulation in which blood from the capillary
beds of the small intestine is brought by
the hepatic portal vein to the capillary
beds of the liver, for further processing.
hepatic portal vein: Large vein carrying
blood from the upper part of the small
intestine, where the blood has absorbed
digested nutrients, to the liver, where it
undergoes further chemical processing
before returning to the heart. Portal
veins carry blood between two capillary
bedsin this case, the capillary beds in
the small intestine and the liverinstead
of between a capillary bed and the heart.
hepatic veins: Two large veins (left and
right) that carry blood from the capillary
beds of the liver to the caudal vena cava
near its entry into the right atrium of the
heart. [485]
hepatoenteric ducts: In birds that lack a
gall bladder, ducts that transport bile from

Haw Hon
the liver to the small intestine. [4121]
herbivores: Organisms that eat primarily
plants. [9123]
Hertz (Hz): A measure of the frequency
of a sound, in cycles (of compression and
thinning of the air) per second. [74]
heterocoelous centrum ends: The saddleshaped, interlocking ends of the centrum
(main body) of avian cervical vertebrae.
Because the anterior end is concave in
a lateral direction and the posterior end
is concave in a dorso-ventral direction,
articulating vertebrae can rotate freely
against one another, allowing the neck
to be highly exible. [415]
heterodactyl foot: Foot arrangement in
which the third and fourth toes point forward and the hallux and second toe point
backward; found in trogons. [121]
heterogametic: Describes individuals
whose two sex chromosomes are different; in birds, the chromosomes are called
ZW and heterogametic individuals are
females, but in mammals, the chromosomes are called XY and heterogametic
individuals are males. Heterogametic
individuals are capable of producing
two different types of gametes (eggs or
sperm)one with one type of sex chromosome and one with the other. For comparison, see homogametic. [4136]
heterozygous: Possessing two alleles that
are different for a given gene; for example,
having one allele for blue eyes and one
allele for brown eyes. (Each allele comes
from a different parent.) For comparison,
see homozygous. [1074]
high-aspect-ratio wings: Wings that are
long, narrow, and unslotted; the length
derives primarily from the lengthened
inner wing (as compared to high-speed
wings, in which the length results primarily from the long outer wing). Highaspect-ratio wings are highly efcient
at producing lift at relatively high ight
speeds, but they are difcult to maneuver, especially during take-offs. They
are found in a few seabirds that are highly
specialized for dynamic soaring over the
ocean, such as albatrosses, shearwaters,
petrels, and some gulls. [542]
high-speed wings: Wings that are tapered,
pointed, and in many cases sweptback,
with unslotted primaries and a high aspect ratio; found in birds such as falcons,
swifts, swallows, terns, ducks, and many
shorebirds. High-speed wings allow
good control and high speeds, but are
energetically expensive to use because
the bird must ap constantly. [538]
hindbrain: The posterior portion of the
brain; it includes the cerebellum and medulla oblongata (brain stem). [436]
hinge post: Central rod of binoculars,
around which the two barrels pivot.

Handbook of Bird Biology


Hoatzin: An odd-looking, leaf-eating
bird that nests in branches overhanging
lakes and slow-moving streams in the
Neotropics; it is the only member of its
order and family (Opisthocomidae). To
avoid predators, nestlings may temporarily leave the nest and clamber about
in trees, aided by small claws on their
wings. [177]
Holarctic: A combination of the Nearctic
and Palearctic zoogeographic regions;
the Holarctic encompasses the Northern
Hemisphere north of the tropics. [170]
homeostatic mechanisms: Mechanisms
that act in various ways to preserve balance. For example, various physiological factors keep the body temperatures of
birds and mammals relatively constant.
And, various ecological factors keep
many populations at a density that uctuates around a fairly stable level. [967]
homeothermic: Describes animals that
are able to keep their internal body temperature constant even when the outside
temperature varies. In birds and mammals, the hypothalamus monitors body
temperature and, if required, triggers responses that warm or cool the individual
to bring it back to normal body temperature. [913]
homing ability: The ability to return to
a specic place. For example, homing
pigeons can return to their lofts when released from great distances away. [580]
homogametic: Describes individuals whose two sex chromosomes are
the same; in birds, the chromosomes
are called ZZ and homogametic individuals are males, but in mammals, the
chromosomes are called XX and the
homogametic individuals are females.
Homogametic individuals are capable
of producing only one type of gamete
(eggs or sperm)one that contains the
same type of sex chromosome as the parent. For comparison, see heterogametic.
homozygous: Possessing two identical
alleles for a given gene; for example, having two alleles for blue eyes. For comparison, see heterozygous. [1074]
honeyeaters: A huge, diverse Australasian family (Meliphagidae, 181 species)
of dull-colored, arboreal songbirds with
medium-length, curved bills. Honeyeaters have a distinctive brush-tipped tongue
for gathering nectar and are important
pollinators, but they also eat insects and
fruits. They feed busily, and often congregate at owering trees. [195]
honeyguides: A nonpasserine family (Indicatoridae, 17 species) of the warmer
parts of the Old World, whose members
are peculiar in their ability to digest wax,
especially beeswax, in addition to their



Hoo Inc

insect prey. Some honeyguides lead humans or other mammals to bee nests, and
then eat the wax that is exposed as the
mammal breaks open the nest. [187]
hooklets: Tiny hooks found on each of the
barbules that branch from the distal side
of each barb of a contour feather; hooklets catch onto the barbules branching
from the proximal side of the next barb
(toward the feather tip), lightly holding
the barbs together to form a smooth, continuous vane. [34]
hormone: Chemical substance secreted
into the blood and thus carried to other
parts of the body, where it may stimulate
or regulate the activities of glands or organs. Hormones are the messengers of
the endocrine system. [469]
hornbills: A family (Bucerotidae, 55 species) of large, toucan-like birds of the Oriental and Afrotropical regions; hornbills
have long tails and huge, down-curved
bills topped by a distinctive casque. They
are famous for the females nesting behaviorshe seals herself inside the nest
cavity with the eggs and young. [191]
horns of the hyoid: Part of the hyoid apparatus; the two bones on each side of
the hyoid that extend backward (caudally) from the tongue bone, running beneath the skull and then curving upward
around the back of the head. [413]
House Finch eye disease: See mycoplasmal conjunctivitis. [971]
hover gleaning: See gleaning. [644]
hovering: A type of ight in which an
aircraft or apping animal remains suspended in air in one place. In birds, hovering usually is achieved by positioning
the body nearly vertically and beating the
wings more or less horizontally, producing just enough forward thrust to balance
wind speed, and just enough lift to compensate for gravity. Hovering is an energetically expensive undertaking. [526]
humeral patagium: A ap of skin extending from the brachium to the trunk of the
bird. [111]
humerus: The bone supporting the brachium (upper arm or wing). [110]
Humphrey-Parkes Nomenclature: The
system of naming plumages and molts
most commonly used by ornithologists
today. In this system, the plumage worn
for the longest time each year (the nonbreeding plumage), usually produced by
a complete molt, is called the birds basic
plumage, and other plumages are called
alternate and supplemental. In addition,
molts are named for the type of plumage
they produce, not the feathers they shed.
In the traditional system, a birds breeding plumage is considered to be the main
plumage. [333, 335]
hybridization: Breeding that occurs be-

tween two individuals of different species. [366]

hydrology: The study of water: its distribution, properties, and patterns of ow
on the earth and in the atmosphere.
hyoid apparatus: A V-shaped unit composed of bones and cartilage, located
between the two halves of the lower
jaw. The hyoid apparatus consists of the
tongue bone (entoglossal) and the horns
of the hyoid, which, respectively, support
the tongue and the muscles that control
tongue movement. [413]
hyperphagia: The dramatic increase in
the amount of food that birds consume
as they prepare to migrate; because these
birds eat more than their body requires in
the short term, they store body fat to be
metabolized for energy during migration.
[563, 924]
hyperthermia: Condition in which the
body temperature rises a few degrees
above normal; if the animal cannot bring
its temperature down, it soon dies. [918]
hypoglossal nerve: The twelfth cranial
nerve; it combines with the vagus (tenth)
and glossopharyngeal (eleventh) cranial
nerves to form a combined trunk that
controls movement of the tongue, larynx,
trachea, and syrinx. [442]
hypothalamus: The ventral portion of the
brain region between the forebrain and
midbrain. The hypothalamus plays a major role in hormonal control of body processes by using special neurosecretory
neurons to control the pituitary gland,
which extends from a stalk below the
hypothalamus. [436, 472]
hypothermia: Condition in which the
body temperature drops below normal.
If the animal cannot bring its temperature
back to near normal, it soon dies. [916]
hypothesis: A tentative explanation for
an observation or phenomenon; hypotheses are usually stated in a way such
that they can be tested via the scientic
method. [65]
Hz: See Hertz. [74]

IBA: Important Bird Area. See Important

Bird Areas Program. [10113]
ileum: The short, nal portion of the small
intestine, running from the jejunum to
the large intestine. [4122]
ilium (plural, ilia): One of the three paired
bones fused to form the pelvic girdle; the
ilium forms the cranial and lateral portion, and in birds is completely fused with
the ischium and the synsacrum. It has a
cup-shaped depression for the attachment of the femur (thigh bone). [424]

Cornell Laboratory of Ornithology

immediate variety: Pattern of singing in

which a bird sings a song type once, then
moves on to a different song type and
sings it once, then goes on to yet another
type, and so on, such that the bird sings
many different songs in its repertoire before ever repeating one. [786]
immigration: The movement of individuals into a population. [949]
imperforate septum: A condition in which
the nasal septum (tissue separating the left
and right nasal cavities) has no opening.
In contrast with a perforate septum (see
separate entry), an imperforate septum appears to decrease an animals sensitivity in
detecting odors. However, it increases the
ability to locate an odors source, because
odors entering one nostril do not mix with
those from the other. [490]
Important Bird Areas (IBA) Program: A
program to identify sites (called IBAs) in
each state that are particularly important
to local populations of breeding, wintering, or migrating birds. The program gathers input from bird watchers and professional ornithologists, designates the sites
it feels are most important, and then encourages public and private stewardship
of the sites. Developed as an international
effort by BirdLife International, the program is conducted in North America by
the National Audubon Society. [10113]
imprinting: A type of early learning in
which a young animal quickly acquires
specic information for certain experiences. For example, after hatching, Greylag Goose goslings rapidly learn to follow
the rst moving object they encounter,
which is usually their parent. [612]
inattentive periods: In avian biology,
blocks of time spent off the nest during
incubation. [899]
inbreeding: The mating of closely related individuals within a population.
The greater the degree of inbreeding, the
greater the chance that deleterious genetic traits will occur in the population.
See inbreeding depression. [966]
inbreeding depression: The reduction in
the average tness of offspring born to
parents that are closely related to each
other, compared to the tness of offspring
born to unrelated parents. Inbreeding depression occurs because closely related
parents share more genes, and thus their
offspring are more likely to receive two
copies (one from each parent) of alleles
that cause deleterious traits or genetic
diseases. For example, in humans, the
genetic disease hemophilia was once
common among some of the inbred royal
families of Eurasia. [966]
incest: A mating between close relatives.
incest taboo: The instinctive avoidance
of breeding with close relatives; it has

undoubtedly evolved in many organisms because it reduces inbreeding depression (see separate entry). Because
determining relatedness can be difcult,
many species seem to avoid incest by not
breeding with any individual with which
they have been in close contact during
their developmental period (which is
usually a sibling or parent). [1075]
incubation: The process by which animals that lay external eggs keep those
eggs at the proper temperature for embryonic development until they hatch
(or the nest fails). Only birds, crocodiles,
pythons, and monotremes (egg-laying
mammals) incubate their eggs. In most
cases, birds sit on their eggs to keep
them warm, but many megapodes bury
themin piles of decaying vegetation,
in long tunnels or broad pits where the
earth is warmed from nearby hot streams
or volcanic cinder elds, or in pits or burrows where bare sand or soil is heated by
the sun. In very hot environments incubation may require cooling the eggs by
shading them, burying them in sand, or
keeping them moist. [893]
incubation patch: See brood patch.
incubation period: The time from the
start of regular, uninterrupted incubation
to hatching. [896]
incubation pouch: A type of brood patch,
found on the breast of some albatrosses,
that is a featherless cavity surrounded by
thick feathers into which the single egg
ts so snugly that it may remain inside
even when the bird stands. [896]
indeterminate layers: Bird species that
will lay additional (replacement) eggs
if one or more eggs are removed from
the clutch during laying. Once they have
begun incubation on a full clutch, however, they cease to lay replacement eggs
if more are removed. For comparison, see
determinate layers. [890]
index: A numerical sequence or other
representation that indicates the relative
level, degree, or amount of something or
some property. For example, an index of
population density, or an index of health.
indicator species: A species that acts
as a canary in a coal mine, because
changes in its population density or distribution provide an early warning that its
habitat is changing in some wayoften
through degradation by human activity.
Birds are good indicators because they
are relatively noticeable and easy to survey, because many bird species are high
in the food chain and thus sensitive to the
bioconcentration (see separate entry) of
toxins, and because many species have
narrow habitat requirements. Also called
biological indicators. [10108]

Inc Int
indigobirds: Members, along with whydahs, of the African genus Vidua (members of this genus are called viduines).
Viduines are colorful seed-eaters that
are brood parasites on other members of
their family, Estrildidae; they are known
for the intricate patterns inside the
mouths of their nestlings, which strikingly resemble those of the nestlings of
their hosts. [8142]
indirect tness: An individuals indirect
tness is the portion of its genes that is
transferred into the next generation (and
eventually beyond) as a result of that
individuals blood relatives producing
offspring. Some of an individuals genes
are propagated when a relative breeds
because the individual shares a percentage of its genes with that relative (the percentage depends on the relatedness), and
because the relative passes 50 percent
of its genes (both those that it shares and
those that it doesnt share) to its offspring.
This contrasts with direct tness, which
is the portion of genes transferred through
the production of an individuals own offspring. [686]
individual recognition: The ability of an
animal to identify other specic individuals. [742]
inertial navigation: Finding your way by
keeping track of all the turns and accelerations you have taken. For example,
logging in your brain the turns and accelerations of an outward trip and then
integrating them to compute a direct
route home. Inertial navigation requires
no outside reference points. [582]
information center hypothesis: The idea
that one benet of colonial breeding is
that individuals who have been unsuccessful at nding food might nd better
feeding grounds by watching at the colony for successful foragers (those who return with food), and then following them
to good hunting spots. [659]
infundibulum: The attened, funnelshaped opening of the oviduct. In birds,
when the ovary releases an egg, the infundibulum moves up to the ovary and
opens, swallowing the egg much like a
snake swallows a rat. [4130]
innate behavior: See instinctive behavior. [65]
innate rhythmicity: The ability to contract without being stimulated by the nervous system, as found in cardiac muscle
cells. [431]
inner ear: Complex structure inside the ear
in the form of a membranous, uid-lled
sac (the membranous labyrinth) oating
inside a bony, uid-lled sac (the bony
labyrinth). The inner ear consists of the
semicircular canals enclosing their ducts
and the vestibule enclosing the utriculus
and the sacculusall of which are con-

Handbook of Bird Biology


cerned with balance and the cochlea

and its enclosed cochlear duct, which are
concerned with hearing. [457]
inner shell membrane: In avian biology,
the membrane just inside an eggs outer
shell membrane; it is thinner and less
coarse than the outer membrane. The
inner shell membrane contains the albumen and adheres tightly to the white of
a hard-boiled egg, making it difcult to
peel. [862, 869]
inner vane: The vane located on the
medial side of a wing or tail feather. On
a wing feather, the inner vane is on the
edge of the wing that trails in ight. In
ying birds, the inner vane is wider than
the outer vane, producing an asymmetry
that aids in ight. [33]
inner wing: The portion of the wing from
the wrist to the shoulder; the secondary
feathers are located on one section of the
inner wing. [523]
innominate arteries: Two large arteries
(left and right) that branch from the aortic
arch and then soon branch again into the
carotid arteries to the head and neck and
the subclavian arteries to the front limbs
(wings). [482, 484]
insect-net theory: A variation on the
cursorial theory of the origin of avian
ight, rst proposed by John Ostrom in
1976. The insect-net theory suggests that
dinosaurian bird ancestors rst evolved
feathers as thermal insulation; and then,
as running dinosaurs clapped the feathered forelimbs together to catch insects,
eventually the motion became apping
ight. [E16]
insemination: The transfer of sperm from
the male into the genital tract of the female; in birds, sperm enter the cloaca.
insertion: On a skeletal muscle, the end
(attachment site) that moves the most
during contraction. [427]
inshore feeders: Birds (or other animals)
that forage close to shore, fairly near their
nesting areas; includes birds such as terns
and many gulls. [668]
insight learning: A modication of behavior that occurs by evaluation of a situation
rather than as a result of previous experience with a particular problem. [614]
instinctive behavior: A behavior that is
triggered in full form, without any learning, the rst time an individual responds
to the releaser; also called innate behavior or instinct. [65]
integumentary system: The skin and
structures that are produced by the skin,
such as (in birds) feathers, color pigments, scales, claws, beak, wattles, and
comb. [43]
intention movements: Movements that
are either incomplete (such as the initial



Int Kin

stages of a behavior) or that indicate what

the performer is about to do. For example,
a bird about to attack may crouch down
and tense its muscles. Many intention
movements have become exaggerated
and incorporated into displays. [630]
interactive playback experiments: Computer-controlled playback experiments in
which researchers can change various aspects of songs played to territorial birds in
the middle of a singing encounter, based
on the subject birds singing behavior. For
example, researchers might choose to
play longer or shorter songs, or songs that
match those of the subject, or they might
respond more slowly or more quickly, or
overlap the subjects singing. [749]
interference: In ecology, a situation in
which one individual actively prevents
other individuals from obtaining a limited resource. It may do so through various
behaviors, such as aggressive interactions that set up dominance hierarchies
or territories. [942]
International Union for the Conservation of Nature (IUCN): International
nongovernmental organization based
in Switzerland and devoted to the conservation of species. In 1963, the IUCN
drafted the original text of the Convention on International Trade in Endangered Species, also known as CITES (see
separate entry), which was nally ratied
in 1975. [1093]
interpupillary distance: The distance between the centers of an individuals two
pupils; important in tting binoculars.
interspecic competition: Direct competition for limited resources among individuals of different species. [956]
intestinal lymph trunk: Lymph vessel that
carries the products of fat digestion from
lymph vessels coming from the small
intestine to the thoracic duct. These digestive products are rst picked up at
the small intestine by lymph capillaries,
which carry them to progressively larger
lymph vessels running along the surface
of blood vessels in the intestinal wall.
These vessels eventually join to form the
intestinal lymph trunk, which carries the
products to one of the two thoracic ducts,
which eventually enter the cranial vena
cava. The reason for this circulatory pattern, which occurs in (at least) all birds
and mammals, is not understood. [489]
intraspecic competition: Competition
(for food, territories, mates, and so on)
that occurs among members of the same
species. [939]
ioras: Small, arboreal songbirds that
search leaves, often in dense foliage, for
insects. The four species form the family
Aegithinidae, endemic to the Oriental
zoogeographic region. [189]

iridescent colors: Structural colors, sometimes brilliant, that shimmer and glitter
because they change in brightness as the
angle of view changes. The colors are produced when light waves reected off thin
lms (in birds, structural layers within attened feather barbules) interfere with one
another, as can be seen in soap bubbles
and most hummingbirds. [354]
iris: The colored part of the eye surrounding the pupil; it is part of the eyes choroid
(middle) layer. The iris contains muscle
bers and controls the diameter of the
pupil and thus the amount of light that
enters the eye. [44]
irruptive migration: Migratory movements that are irregular in time and
space, depending upon factors other
than a change of seasons, such as food
availability. For example, the seeds and
buds eaten by nches such as Pine Siskins
and redpolls uctuate in abundance not
only seasonally but from year to year and
from region to region, so in some years
large numbers of the birds move out of
northern forests to breed, yet in others
they stay put. [555]
ischium (plural, ischia): One of the three
paired bones fused to form the pelvic
girdle; the ischium forms the caudal and
lateral portion. [424]
island endemics: Species that evolved on
an island and are found only upon that
island. [955]
isthmus: The region of the avian oviduct
after the magnum and before the shell
gland; the isthmus secretes the egg and
shell membranes and uid albumen
around the fertilized ovum as the ovum
travels down the oviduct. [4130]
IUCN: See International Union for the
Conservation of Nature. [1093]

jejunum: The long middle portion of the
small intestine, running from the duodenum to the ileum. [4122]
jizz: Birding term for a quick impression
of a birds major features. Jizz harkens
back to the general impression of size
and shape (G. I. S. S.) that British observers used during World War II to distinguish between enemy and friendly
aircraft. [124]
journal: A written record of eld observations of birds, other animals, or natural
phenomena. [247]
jugal arch: Bony rod on each side of the
upper jaw below the palatine (another
set of bony rods). In birds, as the lower
jaw opens it moves the quadrate, which
pushes the palatine and jugal arch forward, pushing on the premaxillary bones
to raise the upper jaw. [412]

Cornell Laboratory of Ornithology

jugular veins: Paired veins (left and right)

carrying blood from the head and neck
region; they merge on each side with the
subclavian and pectoral veins to form
the cranial vena cavae, which return the
blood to the heart. [483]
jugulum: The lower part of the throat of
birds, just below the gular region. [17,
juvenal plumage: Feather coat worn by
juvenile birds after they have molted
their natal down; it consists of the rst
true contour feathers. [329]
juvenile: A young bird. [329]

keel: A midventral ridge of bone that projects outward from the sternum and provides a site for the attachment of the large
pectoral ight muscles. Large, ightless
birds lack a keel, as do the tinamous. Also
called a carina. [423]
keratin: A hard protein that forms scales
and claws and is the primary structural
component of mature feathers. Avian keratin differs from the keratin of all other animals in its amino acid sequence. [328]
kettle: A large aggregation of birds, usually hawks, that are spiraling upward in
a thermal. [539]
keystone species: Species that affect
many other species in their community,
and whose removal would precipitate a
reduction in species diversity or would
produce other signicant changes in
community structure. [9126, 9128]
kHz: See kilohertz. [74]
kidnapping: In avian biology, the aggressive takeover of a brood of young by
adults that are not the parents. [8128]
kidneys: Paired organs of the excretory
system that are irregular in shape and,
in birds, are each composed of three interconnecting lobes; they remove waste
products from the blood, especially nitrogenous wastes, and form a highly concentrated urine that, in birds, is passed to
the cloaca via the ureters. Kidneys also
maintain a balance of salt ions in the
blood. [4125]
kilohertz (kHz): A measure of the frequency of a sound, in thousands of cycles
(of compression and thinning of the air)
per second. One kilohertz equals 1,000
Hertz. Also written kiloHertz. [74]
kingdom: Level of classication of organisms above phylum; similar phyla
are placed within the same kingdom.
All birds are in the kingdom Animalia.
kin selection theory: A line of reasoning
suggesting that the closer the kinship (degree of relatedness) between two animals

of the same species, the greater will be
their tendency to cooperate with one another in various ways. Such cooperation
results from natural selection because
if an animal enhances the survival of a
relative, it also enhances its own indirect
tness (see separate entry); the closer the
relative, the greater the degree to which
its own indirect tness is enhanced.
kiwi: A family (Apterygidae) of three species of grouse-sized, ightless, nocturnal
ratites endemic to New Zealand; kiwis
probe soil with their long beaks, using
their keen sense of smell to locate earthworms. [196]
koilin: See gastric cuticle. [4119]
K-selected species: Species in which
individuals have a life history strategy
that relies on longevity, rather than a
high reproductive rate, to maximize the
number of offspring produced during
their lifetimes. Such species tend to be
large, develop slowly, begin breeding at
a relatively old age, have few young per
cycle (small clutches or litters), breed
infrequently, take care of their young
for extended periods, and have long life
spans. [945]
KT event: The massive, worldwide extinctions 65 million years ago at the end
of the Cretaceous Period and the beginning of the Tertiary Period, which wiped
out dinosaurs as well as many other
plants and animals. The extinctions are
thought to have resulted from worldwide
climatic disturbance caused by a large
meteor colliding with Earth. [E25]

lacrimal gland: One of the two tear

glands on each side of the eye; the lacrimal gland lies in the lower part of the
orbit of the eye and has many ducts entering the space between the lower lid and
the cornea. Its secretions moisten the eye
and nourish the cornea. [446]
laminar ow: A smooth ow of air over
an airfoil. [513]
large intestine: Short, straight tube extending from the small intestine to (in birds) the
cloaca; it holds the intestinal contents while
water is being reabsorbed, and passes the
remainder to the cloaca. [4123]
laryngeal cartilages: Two major cartilages
that make up the sides and oor of the
larynx. Also called the cricoid cartilages.
laryngeal folds: Fleshy folds in the lower
surface of the pharynx that surround
the glottis, the opening to the larynx.
larynx: Structure at the upper end of the
trachea that consists of several cartilages;

Kiw Lim
it acts as a valve, regulating the ow of air
into the trachea. In mammals the larynx
is the voice box, but in birds it produces
no sound, leaving sound production to
the syrinx. [491]
lateral: Toward the side of the body; away
from the midline. [14]
lateral labia: Important sound-producing
membranes in the lateral wall of each half
of the syrinx. During sound production
muscles move the bronchi upward into the
trachea, twisting the third bronchial cartilages so that the lateral labia and medial
labia (in the medial walls) move toward
(and close to) each other and into the path
of air owing out of the respiratory system.
Sound is produced as air rushes between
the lateral and medial labia, causing these
soft tissues to vibrate. [496]
Laurasia: The northern land mass formed
200 million years ago when Pangea split
into two large land masses. It consisted
of present-day North America, Europe,
and Asia. [168]
leafbirds: Eight species of oriole-sized
arboreal songbirds endemic to the Oriental zoogeographic region; leafbirds
are mostly green and yellow, and feed
mainly on insects and fruit. Together with
fairy-bluebirds, they form the family Irenidae. [188]
leaf tossing: A foraging technique in
which a bird tosses aside the leaf litter
(with its beak or feet) to search for food;
used by towhees, turkeys, thrashers, and
many sparrows. [647]
learned behavior: A behavior that requires some amount of previous experiencesuch as exposure to a stimulus
as well as a memory of that experience,
before it is carried out fully. [69]
learning: The modication of a behavior
as a result of experience. [68]
leg spurs: Bony outgrowths near the distal end of the tarsometatarsus, covered by
a pointed horny sheath; used as weapons
by male chickens, peafowl, and many
other pheasant relatives. [426]
lek: 1. A traditional courtship area where
many males of the same species gather
to attract females for mating; each male
spends a large amount of time defending
a small site at which he displays to compete with other males and, in particular,
to earn copulations with sexually receptive females who visit the lek to choose
among the males. In most species, the
top few males secure most of the matings.
The lek contains no nest sites, food, or
other resources useful to nesting females.
[675] 2. The group of males gathered at
the traditional courtship site described in
denition 1. [675]
lek polygyny: Mating system in which
males gather in leks and display to earn

Handbook of Bird Biology


copulations with visiting females. In most

lekking species, the top few males secure
most of the matings, resulting in a high
degree of polygyny. [675]
lens: Spherical or ovoid structure near
the front of the eye; it changes its curvature to sharply focus images from varying
distances on the retina. The lens is crystalline-like and composed of regularly
oriented layers of collagen bers. [447]
Lepidosauromorpha: One of the two major groups of diapsid reptiles; it contains
all snakes and lizards as well as the ancient ichthyosaurs and plesiosaurs. [E8]
lesser coverts: The feathers on the upper
surface of the wing that partly overlie the
median coverts and extend to the marginal coverts. [112, 113]
leukocytes: See white blood cells. Also
spelled leucocytes. [488]
liana: A woody vine. [988]
life history theory: A set of ideas that
attempt to explain the diversity in the
breeding strategies of living things by
looking at why various characteristics
relating to an organisms birth, death,
and reproduction have evolved. These
characteristics, called life history traits
(see separate entry), include such things
as number of offspring, age at rst breeding, and length of the developmental
period. [83]
life history traits: Characteristics of living things that are related to birth, death,
and reproduction. Examples include the
number of offspring, the age at rst breeding, the interval between breeding cycles,
and the chance of surviving to various
ages. Collectively, these characteristics
make up an organisms life history or life
history strategy. [83]
life list: Record of every species seen by a
particular person, noting the date and location of the rst sighting. People may keep life
lists of different groups of organisms, such as
birds, butteries, or reptiles. [254]
lift: Force acting on a moving airfoil (such
as a birds wing) perpendicular to the direction of airow; in a bird that is gliding
or ying horizontally, lift acts upward in
opposition to gravity. [510, 513]
lift-to-drag ratio: The lift produced by a
wing divided by the drag it experiences
while ying. Long, narrow wings have
the greatest lift-to-drag ratios and are the
most efcient. [535]
ligament: Fibrous connective tissue that
connects one bone to another across a
joint. [46]
limiting factor: Something that is present
in (or missing from) a particular environment and as a result prevents a particular
species from living or breeding in that
place. Examples of limiting factors include a large number of predators and



Liv Mat

a low level of a critical resource, such as

food. [910]
liver: The largest internal organ in the
body; in birds the liver has two lobes and
performs a variety of functions, including secreting bile to help emulsify fats for
digestion, storing sugars and fats, forming uric acid, and removing foreign substances from the blood. [4124]
lithornithids: Extinct, ying, chicken-like
palaeognathous birds thought to have
been common in North America and Europe during the early Tertiary; they may
be the ancestral stock that gave rise to
ratites all over the world. [E23]
local population: A population conned
to a small area; for example, all the Blue
Jays in one neighborhood. [949]
lore: Small area between the eye and the
base of the upper beak. [17, 18]
lower beak: The lower half of the beak;
sometimes called the mandible or lower
jaw. [16]
lower critical temperature: The environmental temperature below which
the body of a bird or mammal increases
its metabolic rate and employs physiological responses to warm itself, assuming behavioral responses are no longer
adequate. [916]
lower jaw: See lower beak. [16]
lower lethal temperature: The environmental temperature below which a bird
or mammal cannot keep its body warm
enough to survive. [916]
low pressure system: A weather system
consisting of cold and warm air masses
circulating around an area of low barometric pressure. In the Northern Hemisphere the masses circulate counterclockwise; in the Southern Hemisphere
they move clockwise. [569]
lumbar vertebrae: The vertebrae of the
lower back; in birds they are all fused
with the sacral vertebrae and some of the
thoracic and caudal vertebrae to form the
synsacrum. [415, 418]
lumbosacral enlargement: Swelling
along the spinal cord at the level of the
legs; it is associated with the lumbosacral
plexus. [440]
lumbosacral plexus: Plexus (see separate
entry) along the spinal cord at the level of
the hind limbs; it is associated with the
lumbosacral enlargement of the spinal
cord. [439]
lymph: Fluid carried in lymph vessels;
also see lymphatic capillaries and lymphatic system. [487]
lymphatic capillaries: Tiny vessels that
form a network intertwining with the
capillary beds of the arterial and venous
systems throughout most body tissues.
Some of the tissue uid that has lost its

oxygen and nutrients to the body (tissue) cells and picked up carbon dioxide
and other wastes diffuses into the lymphatic capillaries, and is then termed
lymph. The lymphatic capillaries lead to
progressively larger lymphatic vessels,
which eventually dump their contents
into the venous system. [488]
lymphatic system: Organ systemcomposed of lymphatic vessels, ducts, and
nodesthat gathers tissue uid that has
leaked from the blood capillaries, lters
foreign substances and old or damaged
cells from the uid, and then returns the
uid to the general blood circulation. The
lymphatic system also releases antibodies and transports the products of fat digestion from the intestines to the venous
system, bypassing the liver. [488]
lyrebirds: Two pheasant-sized songbirds
(forming the family Menuridae) of Australian rain forests, named for the elaborate,
harp-shaped tail of the Superb Lyrebird.
They have loud songs and calls and are
fantastic mimics, sometimes clearly reproducing mechanical sounds, such as
logging trucks and chain saws, along
with the songs of other birds. [194]

macroevolution: The evolution of new

species over long periods of time, such
as thousands of years (see microevolution). [137]
magnetic anomaly: A place where the
earths magnetic eld is disturbed, usually by large deposits of iron near the
surface. [598]
magnetic compass: A mechanism by
which birds and some other animals can
use the earths magnetic eld to determine compass direction. [589]
magnetic eld of the earth: The region
around the earth in which objects experience magnetic force (which is a vectora
quantity with both strength and direction).
The eld is created by currents generated
by the constant motion of molten iron in
the earths core. In effect, the earth is a
large magnet with two magnetic poles
(near the North and South Pole). The eld
is stronger closer to the magnetic poles
and weaker toward the equator. [590]
magnetic map: A mechanism by which
birds (and some other animals) may be
able to use the earths magnetic eld as a
map. For example, because the magnetic
eld is strongest toward the poles, a bird
might use the strength of the eld at any
given point to estimate latitude. [597]
magnum: The rst region of the avian
oviduct, after the infundibulum; the magnum is glandular and secretes the rst of
the albumen or white of the egg around

Cornell Laboratory of Ornithology

the fertilized ovum as it travels down the

oviduct. [4130]
malar region: Small area caudal to the
base of the lower beak; also called the
cheek. [17, 18]
malar stripe: A distinctively colored
stripe in the malar region of birds; also
called a mustache stripe or whisker
stripe. [18, 210]
manakins: A Neotropical suboscine family (Pipridae, 44 species) of small, frugivorous birds noted for the stereotyped
courtship displays of the colorful males.
Many manakin species form leks, and
most displays involve intricate maneuvers that may be enhanced by mechanical and/or vocal sounds and plumage
displays. [678]
mandible: In birds, usually refers to the
lower half of the beak; called the lower
beak or lower jaw in the Handbook of Bird
Biology. [411]
mandibular nerve: A division of the trigeminal nerve (the fth cranial nerve)
after it leaves the brain; the mandibular
nerve carries sensory input from the lower beak and corner of the mouth, and motor output from the brain to the muscles
of the lower beak. [441]
manus: The portion of the forelimb distal
to the wrist; also called the hand. The primary feathers attach to the manus. [19]
map and compass model: The theory
(proposed by Gustav Kramer in 1953)
that two things are required to navigate
to a particular destination from an unfamiliar location: (1) a map indicating current location with respect to destination,
and (2) a compass indicating the desired
direction of travel. [584]
marginal coverts: The feathers covering
the upper surface of the wing, from the
leading edge back to the greater coverts,
which they partly overlie. [112, 113]
market hunting: Intensive hunting of species to obtain meat, feathers, hides, or
other body parts highly prized by commercial consumers. [1011]
marine birds: See seabirds. [165]
matched countersinging: Interaction in
which two birds sing back and forth to
one another, each choosing songs that
are identical or similar to those of the
other, or that contain similar phrases.
Not well understood, but thought to be
one way in which birds duel through
song. [736]
matrilineal societies: Groups of animals
in which the females remain in their natal colonies but the males disperse when
they become adults. Thus, within a colony, the females share more genes than do
the males, and are more likely than males
to engage in acts that appear altruistic
toward one another. [656]

maxillary nerve: A division of the trigeminal nerve (the fth cranial nerve) as
it leaves the brain. It carries sensory input
from the skin of the face, upper jaw, upper eyelid, and conjunctiva (tissue covering the eye). [441]
medial: Toward the midline of the body.
medial labia: Important sound-producing membranes in the medial wall
of each half of the syrinx. During sound
production muscles move the bronchi
upward into the trachea, pushing the medial labia and lateral labia (in the lateral
walls) toward (and close to) each other
and into the path of air owing out of the
respiratory system. Sound is produced as
air rushes between the lateral and medial
labia, causing these soft tissues to vibrate.
median: On the midline of the body.
median coverts: The covert feathers lying
between the lesser and greater coverts
on the upper surface of the wing. [112,
medulla oblongata: The most posterior
portion of the brain, also called the brain
stem, where the nuclei of most of the
cranial nerves are located. The medulla
oblongata extends caudally through the
foramen magnum to become the spinal
cord. [437]
megapodes: A family (Megapodiidae, 21
species) of chicken-like terrestrial birds
of the Australasian and Oriental regions.
Megapodes do not use their own body
heat to incubate their eggs; instead, in
many species the males tend the eggs of
several females in huge, warm mounds
of decaying vegetation. Other species
use geothermal heat to warm their eggs.
[193, 670]
melanin: Pigment, usually present as tiny
granules, that produces a range of earthy
colors from dark black, brown, and redbrown to gray, yellow-brown and pale
yellow. Birds can synthesize their own
melanin by oxidizing the amino acid tyrosine. [350]
membranous labyrinth: A system of interconnected, uid-lled canals (containing endolymph) oating in the bony
labyrinth of the inner ear. The labyrinth
forms the cochlear duct, the semicircular ducts, and the utriculus and sacculus.
meninges: General term for the vascularized membranes surrounding the
brain and spinal cord. Specically, these
are the outer brous dura and the inner
arachnoid and pia layers. Meninges provide sustenance and waste removal for
the cells of the brain and spinal cord,
which are not served by the circulatory
system. [436]

Max Moa
mesencephalon: See midbrain. [438]
mesites: A family (Mesitornithidae, 3
species) of rail-like, ground-dwelling
birds endemic to Madagascar; mesites
are about the size of a Mourning Dove.
mesobronchus: The continuation of
each bronchus (right and left) as it enters
the lung and loses its cartilaginous halfrings; also called the primary bronchus.
The mesobronchi gradually decrease
in diameter and branch into secondary
bronchi. Only in birds is the term mesobronchus used as an alternate name for
the primary bronchus. [498]
metabolic water: Water obtained by an
organisms body as a byproduct of the
chemical breakdown of fats, carbohydrates, and proteins within the body.
metabolism: All the chemical processes
that take place in the cells and tissues of
the body. [4144]
metacarpals: Palm bones. In humans
they remain distinct, but in birds, they
are fused with some of the carpals (wrist
bones) to form the large carpometacarpus. [421]
metapopulation: All the individuals of a
species living in an area that contains a
set of subpopulations (local populations)
close enough together so that individuals
from occupied habitat patches can disperse and recolonize patches where the
species has gone extinct. [1079]
metapopulation dynamics: The changes
in local populations found within a metapopulation over time, in which the local
populations (in a habitat patch) may go
extinct and then individuals from within
the metapopulation may recolonize the
patch at a later date, producing a longterm situation in which local populations
wink out and then wink back on.
metapopulation models: See population
viability analysis. [1077]
metatarsals: Instep bones. In humans
they remain distinct, but in birds they
fuse with the distal tarsals (ankle bones)
to form the tarsometatarsus, the long
bone supporting the upper section of the
foot. [114, 425]
metatarsus: See tarsometatarsus. [114]
microevolution: A change in the frequency of a genetically controlled characteristic in a population over a relatively short
period of time (see macroevolution).
midbrain: The middle region of the vertebrate brain; it contains the optic lobes
(which in birds are huge and dominant),
as well as regions for the input and processing of information on hearing and
balance. Also called the mesencephalon.

Handbook of Bird Biology


middle ear: Air-lled chamber between
the eardrum and the cochlea (inner ear);
in birds it houses the columella. [457]
mid-story vegetation: Layer of vegetation
in a forest that consists of trees whose
crowns are below the level of the main
canopy and above the understory vegetation (see separate entry). [993]
migration: The regular movement of all
or part of a population to and from an
area; usually refers to seasonal journeys
to and from breeding grounds or feeding
areas. [552]
migratory program: The genetically
programmed information that guides an
inexperienced bird on its rst migration.
The timing of that migration is controlled
by an internal biological clock that controls circannual rhythms, and the bird
chooses the approximate direction and
distance through its built-in ability to
carry out vector navigation. [579]
migratory restlessness: Nocturnal hopping and uttering during the normal
migratory period, performed by caged
birds that normally are inactive at night.
Researchers often use the degree of this
unrest as an indication of a birds desire to migrate, and the orientation of
the hopping and uttering to indicate
the compass direction in which the bird
would normally be migrating. First called
Zugunruhe by the German scientists who
discovered it. [560]
mimicry: A situation in which one individual or species has evolved or learned
to be similar to another in appearance,
behavior, or sound. For example, Brown
Thrashers and Northern Mockingbirds
imitate the songs of other bird species.
mineral lick: General term for a spot that
animals visit to obtain minerals, either
naturally occurring in the soil or provided
by humans. A clay lick (see separate entry) is one type of mineral lick. [928]
mitigation bank: See wetland mitigation.
units in all cells except bacteria; they
generate most of the cells energy. Mitochondria contain DNA in the form of a
ring. [144]
mnemonic device: A memory aid. In
birding, usually refers to the association
of phrases from human speech with the
songs of particular birds, to help people
remember the songs more easily. [216]
moas: A family (Dinornithidae) of at least
22 species of enormous, ightless ratites
that roamed the open foothills and tussock lands of interior New Zealand. Moas
evolved in the late Tertiary, but were extirpated by the Polynesians approximately



Mob Neo

400 years ago. The largest moa stood 14

feet tall. [196, E24]
mobbing: Behavior in which a number of
birds (often different species) swoop and
dash at a potential predator; they usually
give broad-band, raspy calls (mobbing
calls) that are easy to locate and thus attract additional birds. [226, 651]
model song: A song that a bird listens to
and attempts to duplicate during the process of song learning. [726]
molting: The process of shedding all or
part of the feather coat and replacing it
with new growth. [328] Note: For specic names of molts, see p. 335.
monocular vision: Type of vision that
produces at, two-dimensional images,
in contrast to binocular vision, which
produces three-dimensional images.
Monocular vision results when the eyes
are positioned on the sides of the head
such that an object can be seen by one
eye or the other, but not by both eyes at
the same time. [451]
monogamy: Mating system in which one
male pairs with one female, at least for a
given breeding season. [668]
morph: A set of individuals within a species that are similar to one another in
some genetically determined morphological characteristic, but are distinctly
different from other sets of individuals
within that species. Morphs may differ in
characteristics such as color, body size,
or bill length or shape, but not in characteristics that are related to sex, age, locality, or season. [942]
mortality rate: See death rate. [949]
motor neurons: Nerve cells that convey impulses from the brain and spinal
cord to stimulate a muscle to contract or
permit it to relax, or to cause a gland to
secrete. [433]
mound nest: A nest composed of a pile
of material with an egg chamber in the
middle; in some species the chamber is
entered through a tunnel from the outside. Mound nests are built by many
megapodes, which bury their eggs in a
pile of decomposing organic material;
the Hamerkop of Africa, which builds a
massive mound of sticks; and Monk Parakeets and Palmchats, which build large,
colonial mound nests. [839]
mousebirds: Long-tailed African birds
(family Coliidae, 6 species), about the
size of a Mourning Dove, that climb
through vegetation using their stout,
hooked beaks; frequently mousebirds
perch in a chin-up position, hanging
vertically from a branch. Also known as
colies. [185]
mudnest builders: A family (Grallinidae)
of two striking, black-and-white, robinsized birds named for their large, cup-

shaped, mud nests. The Magpie-lark is

widespread, abundant, and well-known
throughout open areas in much of Australia; the Torrent-lark inhabits fast-owing
streams in the mountainous areas of New
Guinea. [196]
muscle bers: Bundles of long, cylindrical, contractile cells (those that can
contract) that are the functional units of
muscle tissue; they shorten when stimulated by a nerve impulse. [426]
mustache stripe: A distinctively colored
stripe in the malar region of birds; also
called a malar stripe or whisker stripe.
[18, 210]
mutation: A change in the sequence of nitrogenous bases in DNA. Most mutations
have little or no effect on an organism, but
a few result in changes in the structure or
type of proteins produced, thus creating
the genetic diversity upon which natural
selection can act. [136, 141]
mutual displays: Intricate, synchronized
displays or dances performed by members of a mated pair; the displays appear
to stimulate and coordinate breeding
behavior between pair members and to
reafrm the pair bond. Given by many
long-lived birds that mate for life, such
as albatrosses, gannets, grebes, and penguins. [638]
mutualism: An association between two
(or more) organisms in which both (or all)
organisms benet. [97]
mycoplasmal conjunctivitis: A highly
contagious infection of the conjunctiva of the eyes that has spread rapidly
through House Finch populations in eastern North America since its discovery in
Maryland in 1994. The disease also affects Purple Finches, American Goldnches, Evening Grosbeaks, and Pine
Grosbeaks, although to a far lesser extent. Many birds that contract the disease
eventually dieusually from starvation,
predation, exposure to the elements, or
some other factor that results from having impaired eyesight. The disease is
caused by a previously unknown strain
of the bacterium Mycoplasma gallisepticum; other strains had long been known
in poultry. Also called House Finch eye
disease. [971]
myelin: Fatty material forming the nerve
sheath around many axons. The process
by which myelin is deposited in a developing embryo and young animal is called
myelination. [433]
myelination: See myelin. [433]
myelin sheath: The pale, fatty wrapping
that surrounds the axons of many neurons; it insulates the axon, functioning,
in part, like the insulation around household wiring. [433]
myoglobin: A red-pigmented protein

Cornell Laboratory of Ornithology

found in muscle cells. Similar in both

form and function to the hemoglobin
in blood cells, myoglobin binds with
oxygen, storing it until the muscle cell
needs it to release energy. [57]
myopia: Visual condition of an individual in which distant objects blur because images are focussed in front of,
rather than on, the retina; also called
nearsightedness. Some types of birds,
such as penguins, are myopic on land
because their eye is designed for vision
under water. [450]

nape: The back of the neck. [17]

naris (plural, nares): The openings of the
nasal cavity; they are located in the upper
beak, usually near its base. Also called
nostrils. [340]
nasal-frontal hinge: See craniofacial
hinge. [411]
nasal region: Portion of the skull containing the nostrils. [411]
natal dispersal: The movement of a young
animal away from the area in which it was
born and raised. In birds, natal dispersal
usually occurs between edging and the
rst breeding season. [965]
natal down: The soft down feathers covering young birds before they molt into
juvenal plumage. [316]
natural selection: The process by which
evolution occurs: as individuals with
traits that allow them to compete better
for essential resources survive and reproduce better than other members of their
population, they contribute more of their
genes to the next generation, and these
favorable (adaptive) traits increase in the
population. [134]
Nearctic region: Zoogeographic region
including arctic, temperate, and subtropical North America, reaching south
to the northern border of tropical rain
forest in Mexico. In the Handbook of Bird
Biology Greenland is included in the Nearctic, but some sources place it in the
Palearctic. [170, 171]
neighbor recognition: In avian biology,
the ability of territorial birds to identify
their neighborsusually on the basis of
song alone. [743]
Neognathae: One of the two Superorders of ornithurine birds; it contains all
birds with a neognathous palateone
in which the vomer and basipterygoid
process are reduced and a exible joint
exists between the pterygoid and palatine bones (as compared to a palaeognathous palate, which is formed by larger,
more rigid bones). Neognathae includes
all modern birds except ratites, as well as

the extinct Diatryma and dromornithids.
neognathous: See Neognathae. [E23]
neomorphine cuckoo: See cuckoo.
neoteny: An evolutionary phenomenon
in which juvenile traits are retained into
adulthood. [551]
Neotropical migrants: Birds that winter
in the Neotropics but migrate to the Nearctic region to breed. Examples include
many wood-warblers, tanagers, and orioles. [171]
Neotropical region: Zoogeographic region including the West Indies, South
America, and Central America north to
the northern edge of the tropical forests
in Mexico. Also called the Neotropics.
[170, 172]
nerve: A bundle of many nerve cell bers,
surrounded and bound together by connective tissue; one nerve is large enough
to be seen with the naked eye. [432]
nerve cell: See neuron. [432]
nest: In avian biology, a structure built,
excavated, or taken over by a bird, in
which the eggs are laid and remain until
they hatch. In many species, the young
remain in the nest until they are able to
y. In some species, the nest is simply a
scrape or depression on the ground. See
specic nest types, such as cup nest, for
more information. [818]
nest appropriation: Nesting in a nest that
previously was used by another species
or another member of the same speciesusually after the previous breeding
attempt has ended. Nest appropriators
include cavity adopters as well as species that take over open nests, such as
Solitary Sandpipers, Bonapartes Gulls,
House Sparrows, Great Horned Owls,
Little Swifts, and many others. [858]
net primary productivity: See primary
productivity. [987]
neuroglia: Cells that form a supporting,
protective, felt-like bed for neurons and
also provide electrical insulation. Not
nerve cells themselves, they also are
called glia or glial cells. [436]
neuron: A nerve cell. The basic unit of
the nervous system, it consists of a cell
body, axon, and one or more dendrites. It
is capable of generating, conducting, and
receiving nerve impulses. [432]
New World cuckoo: See cuckoo.
New World warblers: A family (Parulidae)
of 115 species of small, insectivorous birds,
many of which are colorful, found in the
New World. Many are Neotropical migrants. Also called wood-warblers. [171]
New Zealand Wrens: A family (Acanthisittidae) of tiny (smaller than a kinglet),

Neo Off
nearly tailless, primarily insectivorous
suboscine birds endemic to New Zealand. One of the four known species, the
Stephens Island Wren, became extinct in
the late 1800s. [196]
NGO: See nongovernmental organization. [10115]
niche: The role played by a particular
species in its environment. Niche includes the many ways in which a species
interacts with its physical and biotic environment, such as what and how it eats,
what temperatures it requires, where it
spends time, when it is active, whether
it disperses seeds or pollinates plants (if
an animal), what animals pollinate it (if a
plant), what preys on it, and so on. Also
called ecological niche. [9102]
niche shifting: Describes a situation in
which a species occupies a different
niche in different communities, depending on which competitors are found in
each. [9103]
nictitating membrane: A thin, translucent
fold of skin that sweeps sideways across
the eye from front to back, moistening
and cleaning the eye and protecting its
surface. Found in many vertebrates, including birds, but not in humans. [17]
night ight calls: Calls given by migrating
birds as they y at night. [744]
nocturnal: Active at night. [931]
node: A clump or mass of tissue, for example lymph nodes [488] or the nodes
in the heart muscle that stimulate the
heartbeat. [479]
nomadic: Pattern of movement in which
individuals are constantly on the move,
showing no tendency to return to previously occupied places. Crossbills and
perhaps Budgerigars may be considered
nomadic. [556]
nonbreeding plumage: See basic plumage. [333]
nongovernmental organization (NGO):
A nonprot organization that is independent from the government. [10115]
noniridescent structural colors: Colors
produced in birds when tiny vacuoles
(pockets) of air within cells in the feather
barbs scatter incoming light. All blues and
whites of birds are noniridescent structural
colors, as are many greens. [355]
North American Waterfowl Management Plan: A 1986 agreement between
the United States and Canada to cooperatively protect waterfowl by jointly
protecting habitat, restoring declining
species, and conducting population
research. Each year biologists make detailed population estimates, and then
use them to help regulate the number
of individuals of each waterfowl species
harvested during the waterfowl hunting
season. [1097]

Handbook of Bird Biology


northeaster: Winter storm that sweeps

into New England across the Atlantic
Ocean from the northeast, the high winds
circulating around a low pressure area.
Often northeasters push pelagic birds
from the open ocean near or over land;
in severe storms, many birds die. Also
called noreaster. [572]
Northern Marine Region: The major faunal region of the seas that includes the
frigid waters of the Arctic south to about
35 degrees north latitude. [199]
northern timberline: See arctic tundra/
coniferous forest ecotone. [9123]
notched tail: See emarginate. [119]
nucleus (plural, nuclei): 1. The membrane-bounded command center of all
cells except bacteria; usually the nucleus
contains the chromosomes. [144] 2.
Clusters of nerve cell bodies within the
central nervous system. Collectively nuclei form the gray matter, found at the
core of the spinal cord and in the outer
areas of the brain. [436]
nuptial plumage: See alternate plumage.

objective lens: The large lens in a pair

of binoculars or a telescope that is farthest from the eye; it receives the image
viewed in the eyepiece. [235]
obligate brood parasites: Bird species that
always lay their eggs in the nests of other
species, leaving the resulting young to be
raised entirely by the host parents. For
comparison, see brood parasite. [8139]
obligate partial migrant: See partial migration. [556]
obligatory annual migration: A type of
migration in which all individuals of a
species migrate to and from a particular
area each year. Usually obligatory migration occurs in species whose breedingarea resources vary greatly from season
to season in a predictable wayfor example, in birds that eat insects or nectar
and that nest at high latitudes where both
insects and nectar become scarce or absent during winter. [554]
occipital condyle: A prominent bump
or peg on the base of the skull, with which
the atlas (the rst cervical vertebra) articulates. Birds have one occipital condyle, but mammals have two. [416]
oculomotor nerve: The third cranial
nerve. It controls eye movement by carrying motor output to the eye muscles; it
also carries motor signals to the eyelid
muscles and the tear gland of the nictitating membrane. [441]
offshore feeders: Birds (or other animals)
that hunt schooling sh far out to sea and



Oil Ost

far from their nests; includes most pelagic

species such as albatrosses and gannets.
Avian offshore feeders are also called pelagic feeders. [668]
Oilbird: A large, nocturnal frugivore of
South America, related to nighthawks;
the Oilbird is the only member of its family, Steatornithidae. Oilbirds use echolocation to reach their nests, which are
located deep within caves. [463]
oil gland: A gland, located at the base
of the tail on the dorsal side of the birds
body, that secretes oils that birds spread
over their feathers during preening. The
oils keep the skin supple and the feathers and scales from becoming brittle,
but they do not appear to waterproof the
feathers. Also called the uropygial gland
or preen gland. [320]
old-growth forests: Virgin (uncut) forests
or forests that have remained uncut for
a very long time and thus contain trees
ranging from hundreds to thousands of
years in age. [1022]
Old World cuckoo: See cuckoo. [8141]
olfaction: The sense of smell. [462]
olfactory: Relating to the sense of smell.
olfactory epithelium: The lining or surface tissue of the nasal cavities; it contains the sensory endings of the olfactory
nerves, which carry input about odors to
the brain. [462]
olfactory lobes: Two lobes at the anterior
ends of the cerebral hemispheres of the
brain that are concerned with the sense of
smell. They are relatively small in birds.
olfactory map hypothesis: The idea that
homing pigeons may learn a gradient odor
map of the vicinity of their home loft by
associating airborne odors with the directions from which winds carry them past
the loft. This gradient odor map would be
based on small but systematic changes in
the intensity or composition of odors over
a large area; as a bird moves in a given direction, particular odors become steadily
stronger or weaker. [593]
olfactory nerve: The rst cranial nerve;
it carries the sensations of smell from the
lining of the nasal cavity to the olfactory
bulb of the brain. [440]
omnivores: Organisms that eat both
plants and animals. [9123]
ology: The study of birds eggs. [880]
operational sex ratio: The ratio of fertilizable females to sexually active males in
a given population. Thus, the ratio measures not just the breeders, but all individuals physiologically and behaviorally
capable of breeding. [980]
operculum: A ap partially covering the
nares; it may help to keep out debris.

Found in some ground-feeding birds

such as starlings, pigeons, and domestic
chickens. [340]
ophthalmic nerve: A division of the trigeminal nerve (the fth cranial nerve) as
it leaves the brain; the ophthalmic carries
sensory input from the nasal cavity (for
nonolfactory nasal sensations), eyeball
(for nonvisual eye sensations), upper eyelid, forehead, and upper beak. In ducks
and geese, it carries sensory input from
the bill tip organ. [441]
opposite birds: A group of small to medium birds that lived in the Cretaceous
period between 65 and 140 million years
ago. They are called opposite birds because their metatarsals (the instep bones
of humans) fuse to form part of the tarsometatarsus from the proximal end to the
distal end, a direction opposite to that
of modern birds. Also called enantiornithines. [E20]
optic chiasma: Site at which the optic tract
coming from the left eye crosses the tract
coming from the right eye just before entering the optic lobes of the brain. [441]
optic lobes: Two large lobes of the midbrain; in birds, they dominate the midbrain and are proportionally larger than
those of mammals. The optic lobes receive the optic tracts from the eyes and
are the sites of much initial processing of
visual information. [437]
optic nerve: See optic tract. [441]
optic tract: Bundle of sensory nerves carrying visual sensations from the retina of
the eye to the optic lobe on the opposite side of the brain. This nerve cable
is considered the second cranial nerve,
although it is really a tract. [441]
optimal foraging: A feeding strategy that
maximizes caloric intake while minimizing the costs of obtaining food, such as
those associated with searching for, capturing, and handling prey. [935]
oral anges: Brightly colored enlargements around the base of the bill in nestlings of many species in which the parents
feed the young. The anges extend from
the corner of the mouth and taper toward
the tip of the bill, and are well-supplied
with tactile nerve endings. Touching a
ange causes the mouth to spring open,
and the colors may help parents to place
the food properly. [343, 8107]
orbit: Cavity in the skull that houses the
eye. The eyes of most birds are so large
that the left and right orbits nearly meet at
the midline of the skull. [411, 413]
order: Level of classication of organisms above family and below class;
similar families are placed in the same
order. The scientic names of bird orders
end in iformes (for example, Passeriformes). [152]

Cornell Laboratory of Ornithology

organ: A group of tissues, often of distinctive character and function, that aggregate to form a discrete structure with
a particular function, such as the heart,
stomach, or lung. [42]
organ system: A group of organs whose
various functions are coordinated to accomplish one or more of the basic functions of life; examples include the digestive system and the respiratory system.
Oriental region: Zoogeographic region
including all of Asia south and east of the
Himalayan Mountains (India and Southeast Asia), as well as southern China and
the islands of Indonesia and the Philippines, east to include the islands of Timor
and Sulawesi. [170, 188]
origin: The end (attachment site) of a skeletal muscle that moves the least during
contraction. [427]
ornithischian dinosaurs: One of the two
major groups of dinosaurs, called ornithischian (bird-hipped) dinosaurs because
their hips supercially resembled those
of modern birds. They were highly specialized herbivores. [E8, E32]
ornithopters: Early ying machines
whose crude wings were lifted by humans apping their arms; they never got
off the ground. [58]
Ornithurae: One of the two major subclasses of birds; it includes all modern
birds as well as the Lithornithids, Ambiortiformes, Hesperornithiformes, and
Ichthyornithiformes. [E20]
ornithurine: Describes birds in the Subclass Ornithurae. [E20]
oscillogram: A visual representation of
a sound, plotted on a graph as relative
loudness (vertical axis) versus time (horizontal axis). The vertical axis actually is
a measure of the increase or decrease in
air pressure (measured in micropascals)
associated with the sound wave, which
determines the loudness. The oscillogram
does not show sound frequency. [77]
oscines: Members of Suborder Passeri,
which is one of the two large suborders
of Order Passeriformes (perching birds);
oscines also are known as songbirds or
true songbirds. Oscines have particularly
complex voice boxes, which allow them
to sing more complex songs than other
birds. [725]
ossication: Hardening or calcication
of soft tissue (such as cartilage or tendon)
into bone or a bone-like material. [46]
Ostrich: The largest living bird and the
only member of its order and family, Struthionidae. This familiar, ightless ratite of
African deserts and savannas is almost
entirely herbivorous. Ostriches breed
communally, with several females laying eggs in the same nest. [183]

otic: Of or relating to the ear. [411]
outbreeding depression: The reduction
in the degree to which local populations
are adapted to their local environments
as a result of individuals dispersing and
mating with individuals in other populations (outbreeding). The reduction occurs
because outbreeding individuals bring
alleles adapted to one local environment
into another, where they may be less advantageous. [966]
outer shell membrane: A loose, brous
membrane that lines the inner surface
of the avian eggshell; it sticks tightly to
the eggshell, helping to hold it together.
[862, 869]
outer vane: The vane located on the side
of a wing or tail feather that is away from
the midline of the bird. On a wing feather, the outer vane is on the edge of the
wing that leads in ight. In ying birds,
the outer vane is narrower than the inner
vane, producing an asymmetry that aids
in ight. [33]
outer wing: The portion of the wing from
the wrist to the wing tip; the primary feathers are located on the outer wing. [523]
ova (singular, ovum): Female reproductive
cells, also called egg cells or eggs, both
before and just after they are fertilized by
a sperm cell. [4128, 4130]
oval: Describes an egg that is shaped like
a chicken egg; also called ovate. [873]
oval window: Former name for the vestibular window. [457]
ovary: The female gonad; it matures
and releases egg cells (ova) periodically
throughout the breeding season in a process called ovulation, and also produces
the sex hormones estrogen, progesterone,
and testosterone. In most birds only the left
ovary is functional, and it enlarges greatly
during the breeding season. [4128]
ovate: See oval. [873]
Ovenbird: A wood-warbler (family Parulidae) that breeds in mature deciduous
forests of the eastern and central United
States and migrates to the Neotropics for
the winter. [835]
ovenbirds: A diverse Neotropical suboscine family (Furnariidae, 240 species)
whose members are especially numerous in temperate South America; ovenbirds are named for the oven-shaped,
clay nests built by some species. [179]
overstory vegetation: The mature trees that
make up the canopy of a forest. [993]
oviduct: The tube that transports the egg
from the ovary to (in birds) the cloaca; it is
suspended from the dorsal body wall by a
curtain-like membrane. After the egg enters
the oviduct as an egg cell (ovum) and is fertilized, different sections of the ovary add
different substances to the ovum to produce

Oti Par
the hard-shelled egg. In most birds, only the
left oviduct is functional. [4131]
oxidation: The process by which a substance is chemically combined with oxygen; also called burning. When body
cells combine oxygen with the products
of food digestion, such as carbohydrates,
the reaction releases energy and two
waste productswater and carbon dioxide. [486]
oxpeckers: Two bluebird-sized species in
the starling family (Sturnidae) that climb
upon large grazing mammals in the African savannas, removing ticks, insects,
and the scabs of skin woundsa behavior that benets both the birds and their
hosts. Also called tickbirds. [187]
oxygenated blood: Blood whose red
blood cells contain high levels of oxygen.
Also called oxygen-rich blood, it is found
in all arteries except the pulmonary artery and its branches. [481]
oxygen-poor blood: See deoxygenated
blood. [481]
oxygen-rich blood: See oxygenated
blood. [481]

Palaeognathae: One of the two Superorders of ornithurine birds; it contains all

birds with a palaeognathous palateone
formed by large, rigid bones (compared
to a neognathous palate in which the
vomer and basipterygoid process are reduced and a exible joint exists between
the pterygoid and palatine bones). Palaeognathae includes all living ratites as well
as the extinct elephantbirds and moas.
[E21, E23]
palaeognathous: See Palaeognathae.
[E21, E23]
palate: The roof of the mouth. [491,
palatine: Bony rod on each side of the
upper jaw above the jugal arch (another
set of bony rods). In birds, as the lower
jaw opens it moves the quadrate, which
pushes the palatine and jugal arch forward, pushing on the premaxillary bones
to raise the upper jaw. [412]
Palearctic region: Zoogeographic region
that consists of most of the large landmass
of Eurasia, as well as northern Africa and
most of the Sahara Desert. In some systems, but not the one used by the Handbook of Bird Biology, Greenland is included
in the Palearctic. [169, 170]
pamprodactyl feet: Foot arrangement
in which all four toes, including the hallux, point forward. Found in some swifts.
pancreas: Digestive organ and endocrine
gland located in the uppermost loop of

Handbook of Bird Biology


the small intestine; the pancreas secretes

digestive juices into the small intestine
and produces hormones (insulin, glucagon, and somatostatin) that regulate
carbohydrate metabolism and blood
sugar levels. [475, 4122]
Pangea: The single land mass that formed
245 million years ago from all the existing continents and persisted throughout
the Triassic period. [168]
Pantanal: An enormous expanse of grasslands extending across parts of Brazil,
Paraguay, and Bolivia that each year is
transformed into a huge marsh by torrential seasonal rains. It hosts one of the
greatest concentrations of wildlife on the
continent of South America, including
numerous waders and other water birds,
as well as the few remaining Hyacinth
Macaws. [173]
pantropical: Distributed throughout the
tropics of the world. [181]
papilla (plural, papillae): A small bump.
1. The bump at the end of each of a
birds deferent ducts where it opens into
the cloaca; also called the papilla of
the deferent duct. When the cloaca is
everted during copulation, the papillae
may slightly enter the females oviduct.
[4128] 2. One of many small bumps
covering the surface of the skin of birds
during embryonic development. Each
papilla consists of a core of dermis and
a covering of epidermis and will eventually form an embryonic feather. Also
called a feather papilla. [326] 3. One
of many spiny-tipped bumps on the undersides of the feet of some birds, such as
Osprey, to aid them in grasping slippery
prey. [345] 4. One of many tiny bumps
in the lining of the shell gland of the oviduct; they secrete albumen and the hard,
calcium-rich outer shell around the fertilized ovum. [4131]
parabronchi: Tiny (microscopic) air tubes
formed by the branching of secondary
bronchi within the avian lung; they form
a network within the lung tissue. Air carrying oxygen passes out of openings in the
thick walls of the parabronchi and into a
network of air spaces surrounded by a
network of blood capillaries. Here, gas
exchange occursoxygen dissolves into
the blood and carbon dioxide and water
vapor move from the blood into the air and
back into the parabronchi, eventually to
be exhaled. The avian system of airow is
much more efcient than that of mammals
(in which air exchange occurs in deadend pockets called alveoli), because in
birds air ows continuously across the
surface of the capillary bed. [498]
paramo: Humid grassland area of the
high Andes in South America above the
tree line. It contains some shrubs and is
dotted with lakes and bogs. [175]



Par Per

parasitic: Describes a relationship between two species or individuals in

which one benets as a result of some
cost to the other. [991]
parasitoids: Animals (usually insects)
that feed in or on a host animal for a long
period of time, consuming most of its
tissues and eventually killing it. Some
parasitoids (such as certain small wasps)
live in the nests of cavity-nesting birds,
feeding on avian ectoparasites and actually beneting the birds. [858]
parasympathetic system: The part of the
autonomic nervous system that acts on
smooth muscle to reduce the heart rate
and to promote feeding, egg laying, and
other peaceful activities, such as digestion. Its nerves originate in the cranial
and sacral region. The other part of the
autonomic nervous system, the sympathetic system, functions under conditions
of stress. [435]
parathyroid glands: Several small endocrine glands located on or caudal to the
thyroid glands; they secrete parathormone, a protein that causes calcium resorption from the bones. [474]
partial migration: Migratory pattern
in which some individuals in a population migrate while others remain as
year-round residents. When the specic
individuals that migrate are determined
genetically, a situation that occurs in environments where the resources always
are sufcient to enable some, but not all,
individuals to overwinter, the migrating birds are called obligate partial migrants; examples include the European
Robin and populations of the Blackcap
in southern Europe. When the number
of birds the environment can support varies from year to year, facultative partial
migration evolves, in which the number
and identity of the individuals migrating
change from year to year in response
to the availability of resources, usually
food. In this case, the individuals that
migrate are not determined genetically.
Facultative partial migration occurs in
the Blue Tit and some North American
chickadees. [556]
partial molt: A type of molt in which only
some of the feathers are replaced. [328]
Passeri: One of the two large suborders
of Order Passeriformes (perching birds);
members are also known as oscines,
songbirds, or true songbirds. Songbirds
have particularly complex voice boxes,
which allow them to sing more complex
songs than other birds. [725]
Passeriformes: See passerines. [166,
passerines: All birds in the Order Passeriformes; also called perching birds.
The order contains approximately 4,600
species, nearly one-half the worlds bird

species, all of which have a foot adapted

for perching on branches or stems. [166,
patagium: A fold of tough skin that extends
from the brachium to the antebrachium;
the patagium connects the shoulder to
the wrist and forms the leading edge of
the inner wing in ight. [111]
patella: The kneecap; an ossication
within a tendon at the lower end of the
femur (thigh bone). The patella glides in
a deep groove and adds stability to the
knee joint. [424]
pecten: Highly vascularized structure of
the choroid layer of the eyes of all birds
and some reptiles; it projects into the vitreous body where the optic nerve exits
the eyeball. The pecten is believed to
nourish the retina and to control the pH
of the vitreous body. [447]
pectinate claw: A modication of the
side of the middle toe claw into a comblike, serrated edge thought to be used
as a preening tool. Found in only a few
birds, such as Barn Owls, nightjars, bitterns, and herons. Also called a feather
comb. [346]
pectoral girdle: The three bones (clavicle, coracoid, and scapula) on each side
of the avian body that form a free-oating support for the wings. [419]
pectoral veins: Paired veins (left and
right) that carry blood from the pectoral
region; they merge with the subclavian
and jugular veins on each side to form the
cranial vena cavae leading to the heart.
pectoralis: Large, powerful ight muscle
of birds that attaches to the sternum. The
pectoralis has two portions: the larger
part pulls the wing down, slows it down
at the end of the upstroke, and pulls the
wing forward; the smaller part pulls the
wing down and back. [56, 57]
peents: Short, nasal vocalizations given
from the ground at twilight by a male
American Woodcock as part of his courtship display. They are similar to the buzzy
call of a Common Nighthawk. [716]
pelagic: Of the ocean; pelagic birds
spend most of their life on the open sea,
feeding at the surface or just below it
and coming to land only to nest. Pelagic
birds include species in the order Procellariiformes, as well as tropicbirds, some
penguins, the boobies and gannets, most
alcids, the skuas and jaegers, the noddies, kittiwakes, Sabines Gull, and some
terns. [198]
pelagic feeders: See offshore feeders.
pellet: A compact ball of indigestible
food, such as bones, fur, feathers, and insect exoskeletons, that is formed by the
gizzard of birds that eat meat or sh

Cornell Laboratory of Ornithology

such as owls, hawks, and kingshers

and is regurgitated through the mouth;
also called a cast. [4119, 4120]
pelvic girdle: The three bones (ilium, ischium, and pubis) on each side of the
body that are partly fused with one another and with the synsacrum in birds to
form a strong but lightweight attachment
site for the muscles of the legs, tail, and
abdomen. They also provide protection
for the abdominal organs. Also called the
pelvis. [424]
pelvis: See pelvic girdle. [424]
pendulous cup nest: A cup nest whose
rim is supported but whose unsupported
belly (nest chamber) hangs from around 4
inches (10 cm) to over 1 yard (1 m) below
the supports. It is generally entered from
the top. Pendulous cup nests are built by
New World orioles, oropendolas, and
caciques, as well as some Old World
weavers, and usually are woven from
plant strips or bers. Shallow pendulous
cup nests grade into pensile cup nests
(see separate entry). [832]
pensile cup nest: A cup nest whose rim is
supported but whose unsupported belly
(nest chamber) hangs below. Pensile cups
are built by New World blackbirds, vireos, and kinglets; by many songbirds in
Australia and Asia; and by a host of orioles and ycatchers and their relatives in
the Old World. Deep pensile cup nests
grade into pendulous cup nests (see separate entry). [830]
perch gleaning: See gleaning. [644]
perching birds: See passerines. [166,
perforate septum: A condition in which
the nasal septum (tissue separating the
left and right nasal cavities) has an opening or is absent. In contrast with having an
imperforate septum (see separate entry),
having a perforate septum appears to increase an animals sensitivity in detecting
odors. However, it decreases the ability
to locate an odors source, because odors
entering one nostril mix with those from
the other. [490]
pericardium: Thin membrane surrounding the heart; its inner lining secretes pericardial uid, which reduces
the friction of the beating heart against
adjacent tissues. [480]
perilymph: The uid that lls the structures that make up the bony outer labyrinth of the inner ear (the cochlea, semicircular canals, and vestibule), in which
oats the membranous labyrinth. [457]
period: A unit of geological time. Periods
are divided into epochs, and successive
periods make up an era. [1113]
peripheral nervous system (PNS): The
portion of the nervous system outside the
brain and spinal cord; it consists of the

cranial nerves as well as the spinal nerves
and their associated ganglia. [440]
pessulus: Cartilaginous structure found
only in songbirds; it sticks up into the
syrinx at the bifurcation of the bronchi.
phalanges (singular, phalanx): The bones
of the ngers and toes. [19, 425]
phallus: See cloacal phallus. [4129]
pharynx: Area, also known as the throat,
that begins at the back of the tongue and
connects the mouth and esophagus, and
where the digestive and respiratory pathways cross one another. The nasal cavities
and auditory tube open into the pharynx.
[491, 4112]
Phasianidae: The pheasant family; it includes pheasants, partridges, ptarmigan,
grouse, guineafowl, turkeys, and Old
World quailas well as the Red Junglefowl (the wild relative of domestic chickens) and the spectacular peafowl. Its 177
species are found throughout most of the
world, but few species inhabit the Neotropics. [189, 190]
philopatry: The tendency of individuals
of certain species to eventually take
up residence (and breed) near the area
where they were born. [965]
photoperiod: The amount of time during
each 24-hour period that an organism is
exposed to light. [563]
photoperiodism: A behavioral or physiological response to day length. [4140]
phylogeny: The evolutionary history of
an organism. [132]
phylum (plural, phyla): Level of classication of organisms above class and
below kingdom; similar classes are
placed within the same phylum. All birds
are in the phylum Chordata. [152]
physiographic regions: Regions of the
world that are biologically distinct based
on climate conditions, topography, soil
types, and plant communities; also called
ecoregions. [1086, 1087]
phytoplankton: Microscopic plants and
algae that oat freely in aquatic environments; phytoplankton make up the
plant portion of plankton. [1103]
pia: The innermost of the three vascularized membranes called meninges
that surround the brain and spinal cord.
The meninges provide sustenance and
waste removal for the cells of the brain
and spinal cord, which are not served by
the circulatory system. [436]
pigeons milk: See crop milk. [4116]
pigments: Colored substances that give
color to the structures, such as skin and
feathers, in which they occur. In theory,
pigments can be extracted from these
structures. [350]

Pes Pop
pineal gland: Endocrine gland located in
the dorsal midbrain; it secretes the hormone melatonin, which plays a role in
regulating daily activity cycles (circadian
rhythms). [471]
pin feathers: Developing feathers that
are still surrounded by a feather sheath.
pinyon-juniper woodland: Ecosystem of
southwestern North America consisting of
park-like stands of small pinyon pines and
junipers growing on hills and mountain
slopes below the montane forests. Five
species of breeding birds are particularly
characteristic to this dry ecosystem: Gray
Flycatcher, Pinyon Jay, Juniper Titmouse,
Bushtit, and Bewicks Wren. [9121]
pipped: Describes an egg about to hatch,
in which the embryo within has punctured a small hole. [8105]
piracy: A foraging technique in which a
bird steals food from another bird; used
by Parasitic Jaegers, frigatebirds, and
some gulls, among others. [648]
pitch: See frequency. [74]
pittas: An Old World family (Pittidae, 31
species) of secretive, stocky suboscines
with long legs and a short tail; they live
on the tropical forest oor, and many
are brightly colored below and cryptic
above. Pittas use their heavy bills to catch
a variety of insects and other small animals, especially snails. [190]
pituitary gland: An endocrine gland attached to the ventral surface of the hypothalamus; it consists of an anterior and a
posterior lobe. The pituitary synthesizes,
stores, and releases a wide variety of hormones. [472]
plankton: Microscopic plants (and algae) termed phytoplankton and microscopic animals (and protozoa) termed
zooplankton that oat freely in aquatic
environments. [1101]
plasma: The uid portion of the blood;
it contains sugars, inorganic salts, and
certain proteins (plasma proteins) found
only in the plasma, and carries numerous
types of dissolved substances. Avian
plasma has a higher sugar, fat, and uric
acid content than the plasma of most
mammals. [486]
platform nest: Nest consisting of a very
shallow depression in the top of a mound
of nest material. Varies in complexity
from the imsy platform of twigs built by
Mourning Doves to the enormous stick
piles amassed by storks or large birds of
prey such as Osprey and eagles. Some
platform nests, such as those of some
terns and rails, oat. [828]
playback experiments: Experiments in
which researchers play songs (through a
speaker) to territorial birds, and then note
their responses. [748]

Handbook of Bird Biology


plexuses: Complex web- or net-like structures along the spinal cord at the level of
the front limbs or wings (brachial plexus)
and hind limbs (lumbosacral plexus);
they are formed by a merging of the large
spinal nerves within each region. [439]
plumage: 1. A birds entire feather coat.
[333] 2. The set of feathers produced by
a particular molt. With this usage, a bird
wears parts of two different plumages after a partial molt. [333]
plunge diving: A foraging technique in
which an airborne bird dives under water to pursue aquatic prey; performed by
birds such as Brown Pelicans, auks, gannets, Osprey, and kingshers. [645]
pneumatic: Describes bones that are
lled with air spaces and may contain
air sacs; nearly all the bones of most birds
are pneumatic. [45]
pneumatic foramen: Opening in the avian humerus (upper wing bone) leading
to the air space within the bone; extensions of the clavicular air sac lead into
the pneumatic foramen and thus connect
with the air space. [4100]
PNS: See peripheral nervous system.
podotheca: The tough skin covering the
tarsus. [120]
polarized light: Light in which the waves
are all oriented in the same plane, or in a
reduced number of planes, compared to
unpolarized light. [594]
polarizing lens or lter: Something that
selectively transmits only those light
waves with certain orientations, creating
a polarized beam of light. [594]
polyandry: Mating system in which one
female mates with several males within
the same breeding season. [677]
polygamy: General term for a mating system in which individuals of one sex (either the males or females) mate with more
than one partner during the same breeding season. Both polygyny and polyandry
are forms of polygamy. [679]
polygyny: Mating system in which one
male mates with several females within
the same breeding season. Polyandrous
species include Spotted Sandpipers, most
jacanas, and certain phalaropes. [668]
polymorphism: A situation in which one
species contains two or more distinct
morphological types (morphs) of individuals, which are determined genetically.
Morphs may differ in physical characteristics such as color, body size, or bill
length or shape, but not in characteristics
that are related to sex, age, locality, or
season. [942]
population: All the individuals of a species that live in the same area. [145,



Pop Pro

population density: The number of individuals per unit area in a given population. [958]
population dynamics: 1. Changes in population size, density, dispersion, and structure over time. [949] 2. The study of how
populations change over time. [949]
population ecology: The study of how
animal populations are related to, and respond to, their environments. It involves
monitoring and studying reproductive
rates, survival rates, movements of individuals and populations, and changes in
population densities over time and from
one area to another. [949]
population structure: The relative proportions of males and females, individuals of different ages, and individuals with
other denable differences in a population. [949]
population viability analysis (PVA): A
computer technique developed by conservation biologists in the 1980s that
simulates the growth of populations over
time. Given numerous demographic variables, such as birth and death rates at various ages and the annual variation in these
rates, PVA can predict the probability that
local populations of different sizes will go
extinct over a designated period of time.
The simplest PVAs are single-population
models, which focus on just one population (of different starting sizes and over
different time periods) and ignore any
dispersal of individuals between populations. More complex PVAs focus at the
metapopulation level (see separate entry);
these use more detailed information, such
as dispersal rates and patterns of behavior
and mortality while moving across inhospitable habitats, to more realistically predict the probability of extinction. [1077]
porphyrins: Pigments producing red,
brown, green, or pink colors in a number of avian orders; they are especially
well known for producing the bright reds
and greens of the turacos. Porphyrins are
complex, nitrogen-containing molecules
related to hemoglobin, that birds and
other organisms synthesize by modifying amino acids. [351]
Porro prism binoculars: Binoculars in
which the eyepieces are closer together
than are the objective lenses. [238]
portal system: A blood circulatory pattern involving portal veins. [485]
portal veins: Veins (such as the hepatic
portal vein and the renal portal vein) that
carry blood between two capillary beds,
rather than connecting a capillary bed
and the heart, as do most veins. Portal
veins allow a second organ to process
the blood before it is returned to the heart
and thus the general circulation. [485]
posterior: Toward the back of an organism, using the earth as a frame of reference.

With birds, technically used only within

the eye and inner ear. But in practice, often
used interchangeably with caudal. [14]
posterior air sacs: General term referring
to the air sacs nearest the birds caudal
endthe posterior thoracic and abdominal air sacs. [4102]
posterior chamber: Space between the
iris and lens of the eye; it is lled with
aqueous uid, which nourishes the eye
and removes wastes. [448]
posterior lobe of the pituitary gland: Part
of the pituitary gland that does not manufacture hormones itself, but instead stores
and releases neurohormones such as mesotocin and arginine vasotocin. [473]
post-hatch brood amalgamation: A broad
term describing a variety of methods by
which young from several broods may
combine into larger groups. [8128]
postpatagium: A tough band of tendinous
tissue running along the edge of the antebrachium and manus that trails in ight.
The postpatagium holds the primary and
secondary feathers rmly in place and
supports each quill. [111]
powder downs: Down feathers that are
never molted but grow continuously,
disintegrating at their tips to produce a
ne powder resembling talcum powder,
which may help to waterproof the feathers. [318]
powered ight: Type of ight in which
the ying object produces thrust. In a
bird, refers to ight powered by apping,
which produces thrust to propel the bird
forward. [536]
precocial: Describes young birds that
hatch in a relatively developed state
downy and with their eyes open. Many
are soon able to walk or swim and even
eat on their own. [8106]
precopulatory displays: Displays given
by either sex immediately before copulation that appear to invite or solicit
copulation. Female birds of many species perform a stereotyped precopulatory
display in which they raise the tail and
hold the wings down or out, quivering
them. [640, 771]
preen gland: See oil gland. [320]
preening: Feather maintenance behavior
in which a bird grasps a feather near its
base, then nibbles along the shaft toward
the tip with a quivering motion; this
cleans and smooths the feather. Many
birds gather on their bill oily secretions
from the oil gland, and then spread them
on their feathers as they preen. [318]
premaxilla: Main bone forming each
side of the upper beak of a bird. The two
halves together are called the premaxillary bones. [411]
prescribed burning: Purposefully starting
res under controlled conditions, at in-

Cornell Laboratory of Ornithology

tervals that mimic the natural re cycle of

the landscape, to maintain certain types
of habitats and the species that depend
on them. [1083]
primary bronchus: See mesobronchus.
primary consumers: Level in a food chain
or web that consists of organismsusually animalsthat directly eat plants
or other producers; most primary consumers are herbivores. [9123]
primary feathers: The ight feathers of
the outer wing; they are attached to the
manus. [111, 110]
primary productivity: Gross primary
productivity is the rate at which green
plants and the few other producers (such
as green algae) in an ecosystem produce
biomass, including the biomass that they
use themselves during respiration. Net
primary productivity is the gross primary
productivity minus the biomass that the
producers use themselves; in other words,
it is the rate at which the producers create
biomass that is available as food for other
organisms. [987]
primary succession: Ecological succession that begins on a substrate, such as
rock, sand, or lava, that has never before
supported a community. Also refers to the
process by which a lake gradually lls in
to form a bog community. See ecological
succession. [9110]
primitive streak: In avian biology, very
early embryonic stage consisting of a
long trough (the streak) with raised
sides that establishes the cranial-caudal,
left-right, and dorsal-ventral body axes.
The primitive streak is formed in the attened mass of cells that lies on the upper
surface of the yolk. [864]
producers: Level in a food chain or web
that consists of green plants and a few
other organisms (such as green algae) that
capture energy from the sun and then introduce the energy into the food chain by
incorporating it into organic compounds,
which the producers synthesize from water and inorganic substances through the
process of photosynthesis. [9123]
productivity: In biology, the total number of grams of living material produced
per square meter per year (g/m2/yr) in a
particular area or habitat. Productivity
includes the increase in size of all living
things, as well as new individuals added
through birth or germination. Productivity
may be measured as gross productivity
or net productivity (see similar denitions
under primary productivity). [992]
projectile vomiting: Vomiting that is
sufciently forceful to propel the regurgitated substance a short distance away.
Birds such as vultures, herons, gulls, and
tubenoses (Fulmars, albatrosses, petrels,
storm-petrels, and diving petrels) use this

technique to ward off predators; Fulmars
can spew an oily, foul-smelling mixture
of esh and uid several feet. [654]
proprioception: A body-parts-position
sense: the subconscious perception of
impulses from the muscles, tendons, and
joints that allows the body to know the
position of its parts, which is crucial for
balance and movement. [433]
proteins: Complex molecules composed
of strings of amino acids; proteins are the
main building blocks of all living organisms and also act as enzymes, assisting
chemical reactions. [141]
protoplasm: The living material composing all organisms. [4144]
proventriculus: The upper part of the birds
two-part stomach; it is elongate and glandular, and secretes mucus, hydrochloric
acid, and an inactive precursor to pepsin,
an enzyme that digests protein. [4118]
proximal: Toward the center of the body
(the elbow is proximal to the ngers) or
toward the origin of a structure (the base
of a featherwhere it is attachedis
proximal to its tip). [14]
proximate: In biology, pertaining to immediate, internal causes; mechanistic.
Proximate questions about a behavior,
for example, look into the internal causes
of the behaviorhow it is carried out. For
comparison, see ultimate. [62]
proximate factors: In biology, the actual
cues that trigger the body to carry out a
particular process or behavior. For example, day length is a proximate factor that
affects the timing of breeding in birds. For
comparison, see ultimate factors. [811]
Prunellidae: A family of 13 sparrowlike birds with slender, pointed bills; it
includes the accentors and Dunnock.
Prunellidae is the only bird family endemic to the Palearctic region. [170]
pruning: A foraging technique in which a
bird bites off and eats plant buds; used by
Ruffed Grouse and other birds. [646]
pseudo-babblers: A family (Pomatostomidae, 5 species) of noisy, busy, social
ground-feeding songbirds with long,
curved beaks and long, towhee-like tails;
they are endemic to the Australasian region. [195]
pseudosuchian thecodont: Term used primarily by Robert Broom in his pseudosuchian thecodont hypothesis to refer to the
thecodonts (see separate entry). [E33]
pseudosuchian thecodont hypothesis (of
bird evolution): Theory that birds evolved
approximately 230 million years ago from
small, arboreal thecodonts; rst proposed
by Robert Broom in 1913. [E8]
pterosaurs: Flying reptiles from the Triassic that radiated and diversied in the
Jurassic and Cretaceous. They had fea-

Pro Rec
tures convergent with those of birds, such
as hollow bones and a slight keel on the
sternum, but their large, membranous
wings were structurally unique: they
were supported by a single, greatly elongated fourth nger and attached to the
side of the body and possibly also to the
hind limb. [E33]
pterylae (singular, pteryla): Areas of a
birds skin where feathers are attached;
also called feather tracts. [32]
pterylosis: The arrangement of feather
tracts and bare patches on a bird. [32]
pubis: One of the three paired bones that
fuse to form the pelvic girdle; the long,
thin pubis runs caudally along the ventral side of the ischium, and is only partly
fused with it. [424]
puddle ducks: See dabbling ducks.
pulmonary arteries: Two large arteries
(right and left) carrying oxygen-poor
blood from the pulmonary trunk (coming from the right ventricle of the heart)
to the lungs; the pulmonary arteries and
their branches are the only arteries that
carry deoxygenated blood. [478]
pulmonary arterioles: Tiny arteries within the lung that supply deoxygenated
blood to the capillaries surrounding the
network of air spaces adjacent to the
parabronchi, where air exchange occurs
in birds. [499]
pulmonary circulation: Portion of the circulatory system carrying blood from the
heart to the lungs and back to the heart.
pulmonary trunk: A large artery that carries oxygen-poor blood from the right
ventricle of the heart and then branches
to the right and left lungs as the right and
left pulmonary arteries. [478]
pulmonary veins: Two large veins (left
and right) carrying oxygen-rich blood
from the lungs to the left atrium of the
heart; the only veins (along with the pulmonary venules) that carry oxygenated
blood. [478]
pulmonary venules: Tiny veins within
the lung that carry oxygenated blood
away from the capillaries surrounding
the network of air spaces adjacent to the
parabronchi, where air exchange occurs.
pupil: The opening in the center of the
iris; it controls the amount of light entering the eye. [447]
pus: The yellowish uid seen in infections; it is a mixture of bacteria, dead
white blood cells, and uid. [488]
PVA: See population viability analysis.
pygostyle: The tail bone of birds; it is
formed by the fusion of the nal few caudal

Handbook of Bird Biology


vertebrae. The pygostyle is shaped like a

plowshare and provides an attachment site
for the ight feathers of the tail. [418]
pylorus: A muscular sphincter (circular
band of muscle) at the lower end of the
stomach; it regulates the passage of food
from the stomach into the small intestine.
pyriform: Pear-shaped. Describes an egg
that is rounded at one end and relatively
pointed at the other; also called conical.

quadrate bone: Bone on each side of the

skull, at the caudal and lower end of the
upper jaw; in birds, it links with the articular bone on each side of the lower
jaw. This linkage forms the joint between
the upper and lower jaws, allowing a bird
to open its mouth widely. [412]
quill: Feather. [111]

race: A population of a species, usually

in a particular geographic area, that is
morphologically distinct from other populations of the same species, but whose
members remain capable of interbreeding with members of those other populations. Also called a subspecies. [155]
rachis: The portion of the feather shaft
above the calamus; it is solid and has a
vane on each side. [33]
radiale: The larger of the two wrist bones
of birds; it consists of fused carpals.
radius: The smaller of the two bones
supporting the antebrachium (forearm).
range: The geographic area within which
a species or population generally remains at a particular time of the year; a
species may have different breeding and
nonbreeding ranges. Also called the geographic range. [219, 949]
raptors: Members of the orders Falconiformes and Strigiformes, which contain
all the diurnal and nocturnal birds of
prey. [166]
ratites: All birds lacking a keel on the
sternum. Includes ightless birdsthe
Ostrich, rheas, Emu, cassowaries, and
kiwisas well as the tinamous, which
are fully capable of ight. [E22]
rattle: A harsh trill. [216]
recolonization: The establishment of a
population of a particular species in an
area where that species formerly occurred, generally through natural dispersal. [1077]



Rec Rol

rectrices (singular, rectrix): The long, stiff

ight feathers of the tail. [112, 113]
recurrent bronchi: Secondary bronchi
(branches off the main air tube, the mesobronchus) within the avian lung that
connect to the air sacs outside the lung.
recurved: Curved upward. Describes
beaks such as that of the American Avocet. [117]
red blood cells: Flattened, elliptical, redcolored cells found in the blood; they
house the iron-containing pigment hemoglobin, which carries large amounts
of oxygen and gives the blood its red
color. The red blood cells of birds have a
nucleus, unlike those of mammals. Also
called erythrocytes. [486]
red bers: A type of muscle ber that
appears red because it contains a large
amount of myoglobin (which carries oxygen) and is permeated by massive capillary beds (containing many red blood
cells, which also carry oxygen). Muscles
with many red bers produce energy aerobically (using oxygen), without building
up the quantities of lactic acid that cause
fatigue, and thus can sustain actions for
long periods of time. Therefore, long-distance iers often have a preponderance
of red bers. Muscles with many red bers often are called dark meat (as in
the drumstick of a turkey). [57]
redirected activities: Actions that are
appropriate for a given situation, but are
directed at an inappropriate subject; often
they appear at times of stress or conicting
motivations. Some have been exaggerated
and incorporated into displays. [633]
reex: A relatively simple and stereotyped response to a stimulus; it may be
either automatic or learned. [434]
refugia: Areas remaining stable when surrounding areas are undergoing change
(such as that caused by glaciation); often
used to describe isolated patches of rain
forest separated by grasslands during glacial periods. [177]
regional population: A population that
inhabits a fairly large area; for example,
all the Canada Geese in the central migratory yway. [949]
reintroduction: The establishment of
individuals of a species (through human
effort) in an area where that species used
to liveusing individuals from a different
area (translocation) or from a captive
breeding program. [1089]
relative abundance (of species): The
number of individuals of each species
living in a particular community or area
compared to the numbers of individuals of
other species in that area. Relative abundance measures how evenly individuals
are distributed among species. [982]

releasers: Specic objects, physical

features, or behaviors that activate (or
release) a specic response in an individual. [65]
relict population: A small, isolated population that is a remnant of a much larger
population that once existed over a wider
geographic area. [957]
remiges (singular, remex): The longest
wing feathers, also called the ight feathers of the wings; they include the primary
and secondary feathers. [111, 113]
renal efferent veins: Veins that collect
blood from the capillary beds of the kidneys and carry it to the caudal vena cava,
which returns it to the heart. [485]
renal portal system: System of blood
ow, found in birds and reptiles but not
in mammals, that collects venous blood
from the lower portion of the digestive
tract via the caudal mesenteric vein, and
conveys it to the venous ring at the kidneys, where some of it passes through
a capillary bed before being conveyed
back to the heart. The reason for this pattern of blood ow is unknown. [485]
renal portal veins: Two veins (left and
right) carrying blood from the caudal
mesenteric vein to the capillary beds of
the two kidneys; part of the renal portal
system of birds. [483]
repertoire: See song repertoire [714,
719, 785] or vocal repertoire. [710]
residents: Individuals that live year round
in a particular area. [554]
resource-defense polygyny: Mating system in which one male may have several
mates, and the females choose their male
partners by evaluating the quality of their
breeding territories; males with better territories attract more mates. This system
may evolve when safe nesting sites or rich
food supplies are more important to the
survival of the young than is male parental care. Found in Red-winged Blackbirds
and a few other species. [673]
reticulate podotheca: A podotheca that
is divided into a network of small, irregular, nonoverlapping plates. [120]
retina: The innermost layer of the eyeball,
upon which images are focused; it is pigmented and contains the light-sensitive
rod cells and cone cells. [449]
retort nest: A globular nest with an entrance tunnel; built out of mud by many
mud-nesting swallows such as Cliff Swallows, and constructed from grass by many
African weavers and New World swifts in
the genus Panyptila. [836]
reverse Porro prism binoculars: Binoculars in which the objective lenses are
closer together than the eyepieces; characteristic of some compact binoculars.

Cornell Laboratory of Ornithology

reverse sexual size dimorphism: A situation in which females are signicantly

larger than males of the same species;
found in many predatory birds such as
raptors, jaegers, skuas, frigatebirds, and
boobies. Because the sexes are different
sizes, they can take different sizes and
thus different types of prey. This may reduce competition between mates, but
why females of these species are larger
than males (rather than the more typical
situation, in which females are smaller)
remains a matter of much speculation.
rhamphotheca: Outer covering (sheath)
of the beak; it grows throughout a birds
life. In most birds it is hard and horny, but
in waterfowl, pigeons, and some shorebirds it is soft and leathery. [339]
rheas: Two species (family Rheidae) of
large, ightless ratites of temperate open
country in South America. [174]
rhomboid sinus: Opening on the dorsal
midline of the lumbosacral enlargement
of the spinal cord; it contains a gelatinous
mass of neuroglial cells rich in glycogen,
known as the glycogen body, whose
function is unknown. The rhomboid sinus and glycogen body are unique to
birds. [439]
rictal bristles: Stiff, hairlike feathers projecting from the base of the beak in birds
that catch insects. They may protect the
face and eyes of some birds that capture
large, scaly insects; they also may help
birds detect movements of prey held in
the beak. [313]
right heart: The right atrium and right
ventricle of the heart; these chambers
contain deoxygenated blood, which
does not mix with the oxygenated blood
of the left heart (left atrium and ventricle).
ritualization: The evolutionary process
by which certain everyday motions become exaggerated, repeated, and stereotyped into displays presenting a clear
message. [632]
rockfowl: Dull-colored African songbirds (family Picathartidae, 2 species)
with colorful, bare heads; they hop along
the rain forest oor in rocky areas and
nest colonially in caves. [185]
rods: One of the two kinds of light-sensitive cells lining the retina of the eye; they
are particularly good at detecting low
light levels (but not in differentiating colors, which is the function of the cones).
When a rod cell is stimulated by light energy, it sends a nerve impulse to the brain
via the optic tract. [448]
rollers: Old World nonpasserines (family
Coraciidae, 12 species) colored in shades
of blue, pink, olive, or chestnut; they are
named for their rolling or rocking dives
during courtship ights. They have long,

slender beaks and forage by perching and
then ying to the ground to catch large
arthropods. [187]
roof prism binoculars: Binoculars with
straight barrels, in which the eyepieces
and objective lenses are directly in line.
rostral: Toward the beak; used for positions on the head and neck. [14]
round window: Former name of the cochlear window. [459]
r-selected species: Species in which individuals have a life history strategy that
emphasizes high rates of reproduction
instead of (and probably at the expense
of) long-term survival. These species are
generally small and tend to develop rapidly, breed at an early age, have many
young per cycle (large litters or clutches),
breed frequently, care for their young
only briey, and have short life spans.
rump: The lower back of the bird, above
the tail coverts; conspicuously colored in
some birds. [16]
runaway selection: One possible explanation for why females of some species
choose the males with the most elaborate
ornaments (such as ornate plumage) to
copulate with. The explanation applies
primarily to females choosing nonpaternal sexual partners, either (1) males for extrapair copulations, or (2) mates in species
in which males do not provide parental
care or other resources for their offspring
(such as territories with food or nesting
sites). The runaway selection hypothesis
suggests that females may choose males
with elaborate ornaments simply because
their sons will then inherit these attributes
and be especially attractive to females,
and thus produce more offspring (of both
sexes); in this way, both the genes for possessing elaborate ornaments and the genes
for choosing male partners with those
ornaments will increase in frequency in
future generations. As each generation of
females chooses the most ornate males
available, the ornaments evolve to be
more and more elaborate, and selection
is said to run away with the traiteven
to the extent that having an elaborate ornament may reduce a males longevity. For
runaway selection to get started, females
must gain some other advantage by choosing males with elaborate ornaments than
simply having attractive sons. Once the
trend is begun, however, the advantage
need no longer exist for runaway selection
to proceed. [684]

sacculus: One of the two membranous

chambers inside the bony vestibule of the
inner ear (the other is the utriculus); these

Roo Sea
chambers contain hair cells and dense
crystals called statoconia, both of which
are embedded in a gelatinous material
that is surrounded by endolymph. The
statoconia and hair cells perceive the position of the head with respect to gravity
(see entry for statoconia). [458]
sacral vertebrae: The vertebrae of the
pelvic region; in birds, they are all fused
with the lumbar vertebrae and with some
of the thoracic and caudal vertebrae to
form the synsacrum. [415, 418]
sagebrush ecosystem: Fairly dry ecosystem found above the lowest elevations
of the Great Basin of North America; it
is a shrubland dominated by sagebrush,
shadscale, and other woody growth 2
to 5 feet (0.6 to 1.5 m) high. Of the few
breeding birds, four species are particularly characteristic: Sage Grouse, Sage
Thrasher, Sage Sparrow, and Brewers
Sparrow. [9121]
sagittal plane: A vertical plane through
the long axis of an organism, extending
from head to tail. It divides the body into
left and right portions. [14]
salivary glands: Glands that secrete saliva, which primarily acts to moisten food
in the mouth. Some birds that obtain their
food from the water, such as Anhingas,
have no salivary glands. The salivary
glands of others, such as certain swifts
and swiftlets, secrete an adhesive substance used in nest building. [4111]
sally gleaning: See gleaning. [644]
sandgrouse: Sixteen bird species that
form the family Pteroclidae and that
inhabit the deserts and dry savannas of
Africa and Eurasia; they are similar in
size and shape to pigeons, but taxonomists are not sure of their relationships to
other families, and thus their order is not
determined. Sandgrouse feed primarily
on dry seeds and often y long distances
to water holes to obtain water. They are
well known for their specialized belly
feathers, which are structurally adapted
to absorb and retain water. Parents (usually the male) soak the belly feathers at a
water hole, and then carry the water back
to their nestlings. [8133, 921]
sap well: A rectangular hole up to a few
inches long, drilled through the bark of
trees and shrubs by a sapsucker. The bird
may return to the well many times to
feed, both on the sap that ows from the
wound and on insects that are attracted
to the owing sap. Other birds and mammals also may feast at the sapsuckers
well. [9128]
saurischian dinosaurs: One of the two
major groups of dinosaurs, the saurischian dinosaurs are also known as
reptile-hipped dinosaurs because of
the structure of their hip joint. They are

Handbook of Bird Biology


composed of two groups, the herbivorous

sauropods and the carnivorous theropods.
[E8, E33]
Sauriurae: One of the two major subclasses of birds; it contains primitive birds
such as Archaeopteryx and the enantiornithines (opposite birds), but no living
bird groups. [E20]
sauriurine: Describes birds in the Subclass Sauriurae. [E20]
scapula: The shoulder blade; one of the
three sets of paired bones that make up
the pectoral girdle. [419]
scapulars: A group of feathers in the
shoulder region. [112, 113]
scavengers: Organisms that eat dead organisms and wastes. [663]
scientic method: A procedure that scientists use to investigate how the world
works. The specic steps typically followed by scientists to investigate aspects
of the world vary among the different scientic disciplines, but in the biological
sciences the scientic method usually
involves the following: asking a question
about the world, formulating the question into a testable hypothesis, designing
a study and collecting unbiased data, analyzing the data to see if the hypothesis is
supported or rejected (this often involves
statistical testingsee separate entry),
and then drawing conclusions. [697]
sclera: Tough, whitish, outer layer of connective tissue surrounding the eyeball.
scleral ossicles: Bony rings supporting
the eyes of all birds, lizards, turtles, and
shes; they are not present in mammals.
scrape: A shallow depression created by
certain types of birds for use as a nest. It
usually contains little or no nest material;
in some species it may be lined with a few
at pebbles. Scrapes are built by many
plovers, terns, skimmers, and penguins.
screamers: A South American family
(Anhimidae, 3 species) of heavy-bodied,
goose-like birds with far-reaching calls.
[174, 422]
scrub desert: Hot, dry ecosystem of the
lowlands and valley oors of southwestern North America; it is composed of
bare ground with widely spaced plants
such as creosote bush, ocotillo, and bur
yucca. The scrub desert ecosystem hosts
a moderate number of breeding birds,
which tend to synchronize their nesting
with periods of rainfall. [9122]
scutellate podotheca: A podotheca that
is broken up into overlapping scales.
seabirds: All birds directly associated
with the open seas and consistently de-



Sea Sif

pendent on the seas for food; also called

marine birds. [165]
search image: A general idea or internal
representation of an object being sought
that allows an animal to nd it easily. If an
animal has a search image for a particular
type of prey, it is able to nd the prey more
efciently than it can nd other prey for
which it has no search image. [925]
secondary bronchi: Branches off the mesobronchus (main air tube) of each avian
lung; some connect to the air sacs and are
called recurrent bronchi, and others divide
to form the tiny parabronchi, the major respiratory units of the avian lung. [498]
secondary cavity nesters: See cavity
adopters. [839]
secondary consumers: Level in a food
chain or web that consists of organismsusually animalsthat directly eat
primary consumers. [9123]
secondary feathers: The ight feathers of
the inner wing; they are attached to the
ulna of the antebrachium. [111, 113]
secondary sex characters: Anatomical
features that distinguish the sexes, but are
not directly involved in the production of
eggs or sperm. [4137]
secondary succession: Ecological succession (see separate entry) that begins
with bare soil or an existing community.
Secretary-bird: A long-legged bird of
prey that stalks the African savannas on
foot for snakes, other small vertebrates,
and large insects. It is the only member of
its family, Sagittariidae. [184]
Section 404, Clean Water Act: Part of
the Clean Water Act (see separate entry)
passed in the United States in 1972; Section 404 strives to protect wetlands by
regulating the lling of, or discharge of
dredged material into, all fresh waters of
the United States, including wetlands.
It allows wetland lling by permit if
applicants show that less damaging alternatives are not feasible, and that the
water will not be degraded signicantly.
Applicants must avoid wetland impacts
when possible, minimize wetland damage, and provide compensation (wetland
mitigationsee separate entry) for all
wetland impacts deemed unavoidable.
seedsnipes: A family (Thinocoridae, 4
species) of ptarmigan-like birds that nest
both in the southern lowlands and the
high mountains of South America. [174]
semicircular canals: Three bony, ringlike canals of the inner ear, arranged at
right angles to one another (one in each
plane of space); they are part of the bony
labyrinth, and thus are lled with perilymph. They enclose the semicircular
ducts, which contain the hair cells that

sense changes in the organisms speed or

direction, thus guiding balance. [458]
semicircular ducts: Three ducts oating
in the perilymph of the semicircular canals of the inner ear, one in each plane
of space; they contain the hair cells that
guide balance by sensing changes in the
organisms speed or direction. They are
part of the membranous labyrinth of the
inner ear and thus are lled with endolymph. [460]
semilunar membrane: Membrane that
extends from the pessulus into the cavity
of the syrinx; it is found only in songbirds.
semilunar valves: Valves, each composed of three cusps (aps), that are located at the beginning of the pulmonary
trunk and the aorta; the semilunar valves
prevent the backow of blood into the
ventricles as the ventricles relax after
each heartbeat. [479]
semiplumes: Feathers that occur in a
continuum of forms between down and
contour feathers. Located at the edges of
the contour feather tracts, semiplumes
provide insulation and help to maintain
the birds streamlined shape. [317]
sensitive period: A relatively brief span of
time early in life during which songbirds
are best able to memorize the details of
songs they hear. [726]
sensory neuron: Nerve cell that conveys
impulses to the spinal cord and brain;
these impulses are interpreted as sensationseither conscious or subconscious.
sentinel system: A system in which group
members take turns watching for danger.
Birds using sentinel systems include
Florida Scrub-Jays and some American
Crows. [656]
septum: Tissue that extends ventrally
from the upper wall of the nasal cavities
and separates the left and right nasal cavities from one another; the septum may
be perforate or imperforate (see separate
entries). [490]
seriemas: A family (Cariamidae, 2 species) of fast, long-legged, grassland birds
of South America; seriemas chase down
their reptile prey on foot. [175]
sex hormones: Hormones produced by
the anterior lobe of the pituitary gland,
the adrenal glands, and (primarily) the
gonads; they regulate the anatomical
structures, physiological processes, and
behaviors that are essential for reproduction. [4137]
sex ratio: The ratio of males to females
(or females to males)usually noted for
a given population. [980]
sexual dimorphism: A situation in which
males and females of the same species differ from each other in size or form. [939]

Cornell Laboratory of Ornithology

sexual selection: Form of natural selection that occurs when individuals differ
in their ability to acquire mates. In sexual
selection the characters that enhance
mating success (either by increasing
an individuals ability to compete with
other members of the same sex and thus
gain mates, or by directly increasing an
individuals attractiveness to the opposite
sex) are selected for, in some cases even
if they decrease individual survival (see
runaway selection). [368, 682]
shaft: The stiff, central rod of a feather,
from which the vanes extend. [33]
shell gland: Region of the avian oviduct
after the isthmus and before the vagina; it
is lined with papillae that secrete uid albumen and the calcium-rich shell around
the fertilized ovum. Shell pigments, if
any, also are added here; the patterns of
pigmentation reecting the speed of the
eggs passage and rotation through the
region. [4131]
shivering: Uncoordinated muscle ber
contraction; a way to produce heat by
muscle contraction without directed
movement. [427]
shock molt: A rare situation in which
a bird sheds many feathers all at once;
it usually is the result of stress, such as
when a bird is grabbed by a predator (tail
feathers may be shed if grabbed by the
tail), handled by humans, or exposed
to violent natural events such as earthquakes and tornados. Also called fright
molt. [338]
Shoebill: A large, stork-like water bird
endemic to Africa; it has a huge, shoeshaped, hooked bill for seizing large sh.
It is the only member of its family, Balaenicipitidae. [184]
shorebirds: Oystercatchers, plovers,
snipes, sandpipers, curlews, phalaropes,
and sheathbills. Ornithologists in Britain
and the British Commonwealth, except
Canada, speak of shorebirds as waders.
shore feeders: Birds (or other animals)
that hunt on shore in the intertidal zone
(between the low and high tide marks),
relatively close to their nests; shore feeders include marine waders, such as the
American Oystercatcher, and many other
shorebirds. [668]
siblicide: The killing of ones sibling; in
birds whose offspring practice siblicide,
usually the stronger, older nestlings kill
the younger, weaker nestlings, either by
direct attack or by pushing them out of
the nest. [687]
sifting: A foraging technique in which a
bird sweeps its partly opened bill from
side to side in water or mud, straining out
small animals and plant material with the
specialized edges of the bill as the water (or mud) drains out. Sifting is used by

Roseate Spoonbills, Northern Shovelers,
and others. [646]
single-population models (in PVA): See
population viability analysis. [1077]
site delity: Loyalty shown by birds or
other organisms to places they previously
occupied; the places may be breeding locations, nonbreeding locations, or stopover points between the two. Also called
site tenacity. [575, 913]
site tenacity: See site delity. [575]
skeletal muscles: Muscles that move the
bones; they constitute the meat of an
animal and are under conscious control.
Also called voluntary muscles. [426]
skulling: A technique by which eld
workers use the degree of skull ossication and pneumatization to determine
the age of live passerines captured in the
fall. [410]
slope soaring: A type of static soaring in
which a bird derives lift from air deected
upward when wind strikes a hill, ridge, or
cliff; or from rising eddies created when
wind spills over a cliff. Slope soaring is
particularly common along seacoasts
among gulls, terns, fulmars, and gannets;
it also is common along mountain ridges, where ravens, crows, and migrating
hawks may soar. [542]
slots: Gaps between the feathers of the
wing tips, created when a bird having
narrow-tipped primaries spreads them
during ight. Slots are most common
in large, soaring birds, such as vultures,
condors, eagles, and certain hawks, because they allow these birds, which have
broad, rounded wing tips, to increase
their lift-to-drag ratio. Slots reduce tip
vortex by turning each primary feather
into an individual, narrow, pointed wing
tip; they also increase lift because each
separated primary feather acts as an individual airfoil. [535]
slotted high-lift wings: Broad wings with
a deep camber and very prominent slotting; the broadness reduces wing loading
and allows low ight speeds, and the slotting increases lift, allowing the birds to
carry heavy prey. Birds such as vultures,
eagles, condors, storks, and some owls
and hawks have this type of wing; they
use it for static soaring. [539]
small intestine: The longest part of the
digestive tract, running from the stomach
to the large intestine. The nal processes
of digestion take place inside the small
intestine, as proteins are broken down
into amino acids, carbohydrates are
converted to simple sugars, and fats are
reduced to glycerol and fatty acids. The
small intestine is longer in birds that feed
on foliage or grain than in those that feed
on fruit or meat, reecting the difculty
of digesting the cellulose in plant material. [4120]

Sin Spe
smooth muscle: Type of muscle found
in the walls of the hollow organs (such
as the stomach or intestines) and in the
walls of blood vessels larger than capillaries, especially arteries and arterioles.
The contraction of smooth muscle is not
controlled consciously. [428]
snood: The limp, red, ngerlike, eshy
structure that projects over the bill from
the forehead of a Wild Turkey. [348]
soaring: In a bird or other animal, ying without apping the wings or limbs,
while gaining altitude or remaining horizontal (as compared to gliding, in which
the animal loses altitude). [536]
social foraging: See cooperative foraging. [664]
Solnhofen limestone: Fine-grained limestone deposits near the village of Solnhofen in Bavaria that, due to meticulous
mining, have produced an array of amazingly well-preserved fossils of plants, invertebrates, shes, and reptiles, as well
as several specimens of Archaeopteryx.
soma: See cell body. [432]
somites: In avian biology, paired segments that appear along the middle
region of the cranial-caudal axis of the
very early embryo; eventually they will
become vertebrae and muscle. [865]
sonagram: A visual representation of
sounds, such as bird song, plotted as a
graph of frequency or pitch (vertical axis)
versus time (horizontal axis). The darkness of the images gives a rough indication of the relative loudness of the different sounds. Also spelled sonogram.
songbirds: Members of Suborder Passeri,
which is one of the two large suborders
of Order Passeriformes (perching birds);
songbirds also are known as oscines or
true songbirds. Songbirds have particularly complex voice boxes, which allow
them to sing more complex songs than
other birds. [725]
song control centers: Groups of nerve
cells within the brains of songbirds that
are involved with both the hearing and
production of songs; they are interconnected with one another in a series of
complex pathways. Individuals that sing
frequently or that have large song repertoires have large song control centers.
The centers shrink in the nonbreeding
seasona time when most birds sing
less often. [739]
song repertoire: All the songs (different
song types) sung by an individual bird; for
comparison, see vocal repertoire. [714,
719, 785]
songs: Loud vocalizations, often delivered from an exposed perch, that are presumed to attract mates or repel territorial

Handbook of Bird Biology


intruders. Strictly speaking, songs are given only by songbirdspasserines in the

suborder Passeri (oscines). In practice,
however, similar vocalizations made by
nonsongbirds (and even other animals)
often are called songs. [714]
Southern Marine Region: The major faunal region of the seas that includes the
frigid waters around Antarctica north to
about 35 degrees south latitude. [199,
southwestern oak woodland: North
American ecosystem that is scattered
on hills and mountain slopes and that
contains open woodlands consisting
primarily of oaks and occasional large
ponderosa pines; southwestern oak
woodland is found in Utah, Nevada, California, New Mexico, Arizona, and parts
of Colorado. Rainfall is low to moderate.
spatial learning: Learning the location of
objects. [612]
speaker replacement experiments: Experiments in which researchers remove
animals (usually male birds) from their
territories, and then replace them with
tape recorders and loudspeakers broadcasting various sounds (the males song,
the song of a different species, background noise with no song, and so on),
depending on the specic experiment.
Then the researchers watch the reaction
of other birds in the area. [769]
speciation: The formation (evolution) of
new species through natural selection.
species: 1. Level of classication below genus; similar species are placed
within the same genus. A species usually is dened as a group of potentially
interbreeding individuals that share distinctive characteristics and are unlikely
to breed with other such groups of individuals. Distinguishing which groups
of organisms constitute species is not always easy, however, and slightly different
criteria may be used by different scientists. [153] 2. The two-word designation
constituting the Latin or scientic name
of a species (see denition 1); the rst
word, capitalized, is the genus and the
second word, lowercase, is the species
name (see denition 3). Both words are
underlined or printed in italics. [147] 3.
The second word in the two-word scientic name of a species (see denition 2).
It is always lowercase, and is underlined
or printed in italics. [147]
species account: A set of written eld
notes on a particular species. [252]
species-area relationship: Typical relationship between the size of an area
and the number of species it supports: as
area size increases, so does the number
of species. This relationship holds (1) for



Spe Stu

patches of the same type of habitat either

in the same or different areas (the larger
the habitat patch, the greater the number
of species it supports), and (2) for areas
of increasing size centered on the same
point (as the boundaries of the area being considered are expanded outward,
the area contains more speciesboth
because the size of a patch of one type of
habitat may increase, and because a larger area may contain more different types
of habitats). The species-area relationship also holds for islands surrounded by
water. [995, 1072]
species population: A population of all
the living individuals of a particular species; also called simply a species. [949]
species richness: The number of species
living in a community (or in a particular
area). The more species, the greater the
species richness. [982]
sperm: See spermatozoa. [4127]
spermatozoa (singular, spermatozoan):
Sperm cells or sperm; each spermatozoan is a single cell composed of a DNAcontaining head and a propulsive tail.
Sperm are short and simple in nonpasserines, and longer and spiral-shaped in
passerines. [4127]
spinal cord: A cable of neurons conducting nerve impulses to and from the
brain; it runs inside the vertebral canal
of the vertebral column, extending from
the medulla oblongata of the brain to the
tail. [438]
spinal nerves: Nerves that arise from the
spinal cord in pairsone on each side
of the cordall along the spinal cord;
each pair carries sensory input and motor
output, and serves a very specic region
of the skin, or very specic muscles or
other organs. [442]
spinous process: A ridge of bone projecting from the dorsal surface of each vertebra; it is particularly well developed in
the thoracic vertebrae. [416]
spotting scope: A medium-range telescope commonly used for bird watching, usually with magnication power
between 15x and 60x. [242]
spread-wing posture: A stance in which
a bird stands motionless with the wings
extended to the side, either to dry the
wings or to absorb sunlight. Adopted by
many large birds, such as cormorants, anhingas, pelicans, storks, and New World
vultures. [322]
spurs: Bony outgrowths from any part of
the skeleton, such as the wing (see wing
spurs). [422]
square tail: Tail shape in which the rectrices are all about the same length. [119]
stabilizing selection: Natural selection
that acts against any change in a current
characteristic. Stabilizing selection oc-

curs when an organism is already very

well adapted to current conditions, and
it tends to keep the species at an optimal
middle point. [137]
stable age distribution: Property of some
populations such that the relative proportions of individuals of different ages are the
same each year, when the same time of year
(such as the start of the breeding season) is
compared. Note, however, that in a stable
age distribution these proportions still
change predictably throughout each year:
in most birds, for example, the proportion
of very young individuals increases in late
spring as eggs hatch, and then gradually
decreases as many juveniles die before
reaching one year of age. [980]
star compass: The mechanism by which
nocturnally migrating birds are able to
use the star patterns surrounding the
North Star (in the Northern Hemisphere)
to determine which way is north (and thus
the other compass directions as well). A
young bird initially learns how to use the
star patterns by observing their rotation
and using the point at the center of their
rotation (the North Star, Polaris) as the
direction to North; but once a bird learns
the star patterns, it can nd north even in
a stationary planetarium. [586]
statant cup nest: Cup nest constructed
on top of a hard physical support or supports, including the ground, building
ledges, or tree branches; built by American Robins, Horned Larks, and many
other birds. [830]
static pressure: The force, produced by
random motion of molecules, that air
exerts uniformly in all directions. When
you squeeze a balloon, you feel the static
pressure from the air inside. [511]
static soaring: A type of soaring (ying
without apping or losing altitude) in
which birds take advantage of the energy
in rising air masses (either in thermals,
called thermal soaring, or along hills or
ridges, called slope soaring) to obtain lift
with little or no energy expenditure on
their part. Performed by birds with slotted
high-lift wings. [539]
statistical testing: Mathematical procedures used by scientists to determine
whether a set of observations or the results
of an experiment either (1) support their
hypothesis, or (2) could result from chance
alone. Most statistical tests generate a
number that indicates the probability that
the observations or results could have
occurred from chance alone. The lower
the probability, the greater the statistical
signicance of the data. Many scientists
choose a 1 in 20 probability (.05) as the
cut-off for signicance: if the probability is
greater than 1 in 20for example, 2 in 20
(.10)then the hypothesis is rejected, but
if the probability is equal to or smaller than

Cornell Laboratory of Ornithology

1 in 20for example, 1 in 100 (.01)the

hypothesis is accepted. In the biomedical
sciences, where the risk of making a mistake may carry serious consequences, acceptable levels of signicance often are
set much lower. [697]
statoacoustic nerve: See vestibulocochlear nerve. [442]
statoconia: Dense crystals of calcium
carbonate embedded, along with sensory
hair cells, in a gelatinous material inside
the utriculus and sacculus of the inner
ear. As the heads orientation changes
with respect to gravity, the crystals settle
against different hair cells, stimulating
them; the brain determines the heads
position by detecting which hair cells
are stimulated. [461]
stepping stones: In conservation biology,
small pieces of habitat, suitable for a particular species, that are located between
larger habitat preserves; they allow individuals to move between the larger preserves. [1080]
sternal rib: The ventral portion of each
thoracic rib; it articulates with the sternum ventrally, and with the vertebral rib
dorsally. [418]
sternum: The breastbone; it provides an
attachment site for the ribs and pectoral
muscles. In all ying birds except tinamous, the sternum has a keel (carina).
stochastic: In biology, describes events
whose occurrence and/or outcomes are
the result of random (or at least unpredictable) factors. For comparison, see
deterministic. [977, 1067]
stomach: Saclike, expandable digestive
organ between the esophagus and the
small intestine; in birds, the stomach
consists of two parts: the proventriculus
(upper) and gizzard (lower). [4118]
stooping: A foraging technique used by
falcons in which a bird drops through the
air at great speed in pursuit of a ying bird
or insect. [645]
structural colors: Colors not produced
by pigments, but instead by the reection
of certain wavelengths of light off the actual physical structure of the object (such
as a feather of a bird or a scale of a buttery wing). [350]
studbook: A collection of information
on the individuals of a particular species. Key to a studbook is data on the
relatedness among individualsfor
example, which individuals are the parents, siblings, and offspring of a given
individual. But studbooks may contain a
variety of other information as well, such
as data on each individuals disease history, birth location and date, and location
in the wild or in captivity. Studbooks are
used primarily by researchers who are

breeding endangered or declining species; the studbook information helps the
researchers to pair individuals with the
partners to whom they are least related,
thus promoting and preserving as much
genetic diversity as possible. [1076]
subadult: A bird that has not yet reached
maturity; immature. [330]
subadult plumage: Any of the plumages
worn by young birds before they reach
their denitive plumages (those of a mature bird). [330]
subclavian arteries: Two arteries (left and
right) that carry blood to the front limbs
(wings). [482, 484]
subclavian veins: Paired veins (left and
right) that carry blood from the front limbs
(wings); on each side, they merge with
the jugular and pectoral veins to form the
cranial vena cava, which returns blood to
the heart. [483]
subelliptical: Describes an egg that is
longer than an elliptical egg, with a slight
bulge to the sides. [873]
suboscines: Members of Suborder
Tyranni, one of the two large suborders
of Order Passeriformes (perching birds).
The suboscines have less complex voice
boxes than members of the other passerine order, Passeri, and thus must sing
simpler songs. Includes birds such as tyrant ycatchers, antbirds, manakins, and
cotingas. [725]
subsong: Practice singing (somewhat like
human infant babbling) that begins shortly after songbirds leave the nest. Subsong
usually is quiet, garbled, and rambling
compared to adult song; it contains some
of the same elements, but they are strung
together in odd ways. It gradually comes
to resemble adult song. [727]
subspecies: A subset of a species, usually in a particular geographic area, that
contains individuals that are morphologically distinct from other individuals
of the same species, but are still capable
of interbreeding with those other individuals. Also called a race. [155]
sugarbirds: A family (Promeropidae,
2 species) of long-billed, long-tailed,
nectar feeders that specialize on Protea plants on the mountainous slopes of
South Africa. [185]
sunbirds: A family (Nectariniidae, 124
species) of tiny birds with long, slender,
down-curved bills; they feed on nectar
and insects and serve as pollinators of
owers. Sunbirds are similar to hummingbirds, but do not hover while feeding. Many males are brilliantly colored.
Most species inhabit Africa, but some are
found in tropical parts of the other Old
World regions. [186]
sun compass: A mechanism by which birds
and some other animals use the position

Sub Syn
of the sun in the sky to indicate compass
direction. To do this, they must be able to
compensate for the changing position of
the sun during the day. [584]
sunning: Exposing the plumage to the
sun, usually by spreading the wings and/
or tail (often against the ground), ufng
the feathers, and remaining motionless;
sunning may aid Vitamin D production
and remove ectoparasites, as well as
warm the bird. [322]
superciliary line: See eyebrow stripe.
[17, 210]
supplemental plumage: In the Humphrey-Parkes system of nomenclature, a
plumage that may occur in addition to the
basic and alternate plumages; it is found
in just a few species, such as ptarmigans
and certain buntings. [333]
supracoracoideus: The powerful ight
muscle (also called the supracoracoid
muscle) that raises a birds wing; it also
slows down the wing at the end of the
downstroke and accelerates it at the beginning of the upstroke. [56]
supracoracoid foramen: See foramen
triosseum. [420, 56]
surface diving: Diving under water from a
swimming position on the waters surface;
performed by birds such as diving ducks,
grebes, cormorants, and loons. [645]
surface-to-volume ratio: An individuals
surface area (through which heat is lost)
divided by its volume (which generates
heat); the larger the ratio, the faster an individual will lose heat. Smaller birds have
larger ratios than larger birds, and thus to
compensate they must have higher metabolic rates and must consume relatively
more food. Also called surface area-tovolume ratio. [311, 4146]
surrogate: In conservation biology, a
stand-in for another individual or species. For example, researchers practiced
releasing techniques for the highly endangered California Condor by rst using
the biologically similar but less endangered Andean Condor as a surrogate.
survival rate: 1. The proportion of individuals in a population that survive for
a particular interval of timeusually a
year. [964] 2. The chance that a particular individual will survive a given
period of time, usually one year. For example, an adult Royal Albatross has an
annual survival rate of 95 percent. [83]
survivorship: The proportion of individuals from a cohort (a group containing individuals born during the same period of
time) that survive to a given age. [85]
survivorship curve: A graph of the proportions of individuals in a cohort (a
group containing individuals born during
the same period of time) that remain alive

Handbook of Bird Biology


at different ages. The survivorship curve

shows the chance of mortality at different
stages in an organisms lifetime; the basic
shape of the curve for a particular species
or group of species illustrates the general
pattern of mortality. [85]
sutures: Boundary lines along the junction between two bones; sutures are visible in the skull of a young bird. [410]
sweeping: A foraging technique in which
a bird pursues insects in ight, capturing them in midair in its mouth; used by
swifts, nighthawks, and many swallows,
among others. [644]
syllable: In a bird song, a note or cluster
of notes that forms a unit that is repeated.
symbiosis: An ecological relationship
between organisms of two different species that live in close association with one
another; types of symbioses include mutualism, commensalism, and parasitism
(see separate entries for each). [9132]
sympathetic ganglion: A clump of cell
bodies of sympathetic neurons that arise
in the spinal cord. A chain of sympathetic
ganglia runs the length of the spinal cord
near its ventral surface. [444]
sympathetic system: The part of the autonomic nervous system that functions
under conditions of stress; the sympathetic system prepares the body for
ght or ight by increasing the heart
rate, breathing rate, and blood pressure.
It consists of nerves that leave the spinal
cord from the thoracic and lumbar regions. [444]
synapse: A small gap between nerve
cells, across which nerve impulses travel.
synchronous hatching: Pattern of hatching in which all the eggs of a single clutch
hatch at about the same time (on the same
day), resulting in a brood of young all the
same size and age. This hatching pattern
occurs when incubation is delayed until
the last egg is laid. Because development
of laid eggs does not begin until they are
warmed, all the embryos begin to develop at the same time and are ready to
hatch at the same time. [688]
syndactyl feet: Foot arrangement in
which the hallux points backward and
toes two, three, and four point forward,
with toes two and three (the inner and
middle toes) fused for much of their
length. Found in many kingshers and
hornbills. [121]
synsacrum: The segment of the vertebral
column of birds that is formed by the fusion of some thoracic vertebrae with all of
the lumbar, all of the sacral, and the rst
few caudal vertebrae; the synsacrum is
in turn fused on either side with the ilium
bones of the pelvis. [418, 424]


syringeal muscles: The muscles of the

syrinx; they allow the syrinx to change
shape and thus to produce different types
of sounds. [493]
syrinx: The sound-producing organ of
birds; located at the point where the trachea divides to form the two bronchi.
[493, 738]
systemic circulation: Portion of the circulatory system that carries blood from
the heart to the body tissues (excluding
the lungs) and back to the heart. [477]


Syr Tip

taiga: See boreal forest. [9116]

tail bone: See pygostyle. [418]
tapaculos: A Neotropical suboscine family (Rhino-cryptidae, 52 species) related
to antbirds; tapaculos have cocked tails
and frequent more southerly, open, and
dry areas than do antbirds. [179]
tarsals: Ankle bones. In humans, they
remain distinct; in birds, the proximal
tarsals fuse with the tibia to form the tibiotarsus, and the distal tarsals fuse with
the metatarsals (instep bones) to form the
tarsometatarsus. [114, 425]
tarsometatarsus: The long bone supporting the upper section of the bird foot;
the tarsometatarsus is formed by a fusion
of the distal tarsals (ankle bones) and the
metatarsals (instep bones). Also called
simply the metatarsus. [114]
tarsus: The upper section of the avian foot,
between the heel and the toes. [115]
taste buds: Simple structures, usually
embedded in the epithelium of the oral
cavity and the tongue, that are the receptors for taste sensations in all vertebrates.
Humans have numerous taste buds located on the tongue, but birds have few taste
buds, which are located primarily on the
roof of the mouth or deep in the oral cavity, with none on the tongue. [465]
taxonomy: The classication of organismsassigning names and relationships. [132]
tectorial membrane: The upper membrane of the cochlear duct in the inner
ear. Sound waves set the lower membranecalled the basilar papilla in
birdsinto motion, moving its hair cells
against the tectorial membrane and triggering nerve impulses in the hair cells,
which are sent to the brain in the process
of sound perception. [459]
telencephalon: The anterior portion of
the forebrain; it contains the olfactory
lobes. [436]
temperate: Moderate; describes a climate free from extreme heat and cold,
but experiencing some of both; generally
found in the middle latitudes. [169]

temporal fovea: Areain the posterior

quadrant of the retina of hawks and other fast-ying diurnal avian predators
where the cones are most concentrated
and the neural layer (the nerves from the
rods and cones, which overlie the rods
and cones and block some light) is the
thinnest, and thus vision is the sharpest.
The temporal foveae provide sharp binocular views of the area in front of the
bird. [449, 453]
tendon: A band of brous connective tissue that attaches muscles to bones, and/
or binds many muscle bers together to
form a skeletal muscle. [46]
teratorns: Giant, soaring, scavenging
birds of the New World (subfamily Teratornithinae) whose fossils date from the
late Tertiary period; teratorns became
extinct around 10,000 years ago when
many large mammals were wiped out.
The largest species had a wingspan approaching 19.4 feet (5.9 m). [108]
termitary: An active termite nest. [819]
territory: A defended area. [622]
tertiary consumers: Level in a food chain
or web that consists of organismsusually animalsthat directly eat secondary
consumers. [9123]
testes (singular, testis): The male gonads;
one testis lies at the cranial end of each
kidney. The testes produce sperm as well
as the male sex hormone, testosterone,
and in some species, a female sex hormone, estrogen. [4127]
thecodonts: A diverse group of reptiles
from the early Mesozoic, all of which have
a diapsid skull, teeth set in sockets, and an
antorbital fenestra (an opening on each
side of the skull in front of the eye socket).
Also known as basal archosaurs or pseudosuchian thecodonts. [E8, E33]
thermal: A rising column or large bubble
of warm air that results from differential
heating of land surfaces by the sun; thermals develop over areas that are darker
than surrounding areas, and over southfacing slopes. [539]
thermal soaring: A type of static soaring
(see separate entry) in which birds use the
rising air in thermals (rising columns of
warm air) to propel themselves upward,
circling higher and higher with little energy expenditure of their own. Once high
in the air, they can glide out of the thermal
and across the countryside in whatever
direction they wish to travel. [539]
thigh: The upper leg, supported by the
femur. [114]
thoracic air sacs: Two pairs of air sacs
in the chest region of birds, one pair (a
cranial and caudal sac) located on each
side of the body. [4101]
thoracic cage: The rib cage; it consists of
the ribs connected to the thoracic verte-

Cornell Laboratory of Ornithology

brae above and to the sternum (breastbone) below. It forms a exible but strong
enclosure for the heart, liver, and lungs,
as well as for the thoracic air sacs of birds.
thoracic ducts: Paired lymph ducts that
collect the products of fat digestion from
the intestinal lymph trunk (coming from
the small intestine; see separate entry);
the thoracic ducts run along the surface
of the aorta and eventually deliver their
contents to the venous system at the cranial vena cavae just before these large
veins enter the heart. [489]
thoracic vertebrae: The vertebrae of
the thorax (chest) region; they articulate
with the ribs and thus form part of the rib
cage. In birds, some thoracic vertebrae
are fused with lumbar, sacral, and some
caudal vertebrae to form the synsacrum.
[415, 418]
thrombocytes: Nucleated blood cells
that resemble red blood cells, but are
more dense and complex and are highly
specialized to carry out blood clotting.
They are not present in mammals, which
instead have platelets (not found in birds)
to carry out blood clotting. [488]
thrust: In a bird, the portion of the force
generated by the apping wings that propels the bird forward (in the direction the
bird is moving, which is always the direction opposite to drag). [518]
thyroid glands: Paired endocrine glands
at the base of the neck. Under the control of the anterior pituitary, the thyroid
glands secrete the hormones thyroxin
and triiodothyronine, which regulate the
annual increase in gonad size, sperm and
egg production, the growth and pigmentation of feathers, and molting. [474]
tibia: In humans, the larger of the two
lower leg bones; in birds, the tibia fuses
with several tarsal (ankle) bones to form
the tibiotarsus, which supports the crus
(lower leg). [114]
tibiotarsus: Bone supporting the lower leg
(crus) of birds; the tibiotarsus is formed by
the fusion of the tibia with the proximal
tarsal (ankle) bones. [114, 425]
tickbirds: See oxpeckers. [187]
time compensation: In the course of
orientation or navigation, making allowances for the changes in the position of
the sun (or other celestial bodies) in the
sky throughout the day. [584]
tinamous: Primitive, grouse-like birds
(family Tinamidae, 46 species) whose eerie calls haunt both forests and pampas of
South and Central America; tinamous are
the only ratites capable of ight. [724]
tip vortex: A type of turbulence created
at the tip of a wing; it consists of air currents spiraling off the wing tip behind a
bird or other ying object. These eddies

sometimes may be seen as trails of white
behind the wing tips of large airplanes as
they take off or land. [534]
tissue: An aggregation of cells with related
and often very similar characteristics; examples include muscle tissue and bone.
tissue uid: Blood plasma that has moved
out of blood cells at the capillary beds,
and is in the minute spaces between the
body cells. Oxygen and nutrients diffuse
out of it to the body cells, and carbon dioxide and other wastes diffuse in. Tissue
uid eventually returns to the blood system, either by diffusing directly back into
a blood capillary near the venous side, or
by diffusing into a lymphatic capillary (in
which it is called lymph), which carries
it to larger lymph vessels and eventually
into the venous system. [487]
tongue bone: See entoglossal. [414]
torpor: A profound state of sleep in
which the body temperature drops and
consequently all metabolic processes
and stimulus-reaction processes slow
down. Used by swifts, hummingbirds,
nighthawks, and some other birds to
conserve energy when food resources are
unavailable. Birds may enter torpor on a
nightly basis, or for extended periods up
to an entire season of cold weather. Also
called hibernation. [4154]
trachea: The windpipe; the tube conducting air from the larynx to the lungs.
tracheal bulla: An expanded sac on one
side of the lower end of the trachea in
males of many duck species; it is thought
to modify the sounds produced by the
syrinx. [492]
track: A ying birds actual direction of
movement with respect to the ground; due
to crosswinds, a birds track may not be the
same as its heading. Also applies to aircraft
and other ying organisms. [568]
tracts: Bundles of axons and their myelin sheaths within the central nervous
system; collectively, tracts form the white
matter, found in the outer portions of the
spinal cord and the inner areas of the
brain. [436]
transequatorial migrants: Birds that migrate back and forth across the equator to
take advantage of spring and summer in
both hemispheres. [1103]
transitive inference: The ability to understand and infer linear relationships
among individuals. For example, the
ability to understand that if A is dominant to B, and B is dominant to me, then
A is dominant to me. [618]
translocation: Establishing individuals of
a species in an area in which that species
formerly lived by importing individuals
from a different area. [1089]

Tis Tym
transverse plane: A vertical plane through
an organism, dividing the body into cranial and caudal portions. [14]
trees-down theory (of the origin of avian
ight): See arboreal theory. [E18]
trial-and-error learning: Learning to associate ones own behaviors with either
a reward or punishment; behaviors that
result in reward are repeated, and those
that result in punishment are abandoned.
trigeminal nerve: The fth cranial nerve;
it divides into the ophthalmic, maxillary,
and mandibular nerves (see separate entries) after exiting the brain. [441]
trill: A phrase or song that consists of a
single note or cluster of notes (a syllable)
that is repeated many times in rapid succession. [216]
triosseal canal: See foramen triosseum.
[420, 56]
trochlear nerve: The fourth cranial nerve;
it carries motor output from the brain to a
single eye muscle. [441]
trophic: Pertaining to feeding. [9123]
trophic levels: The different levels of food
production or consumption within a food
chain or web; for example, producers,
primary consumers, secondary consumers, and decomposers. [9123]
scheme of a community that is based on
the feeding relationships between organisms: species are assigned to different trophic levels according to what they eat
and what eats them (see trophic levels).
Tropical Marine region: The major faunal
region of the seas that includes the warm
equatorial waters between the Subtropical Convergences of both hemispheres.
[199, 1101]
true navigation: The ability to take the
proper course toward a specic goal,
even from completely unfamiliar sites at
vast distances from known areas. [583]
trumpeters: A family (Psophiidae, 3 species) of large, hump-backed, chickenlike birds that inhabit the rain forests and
oodplains of Amazonia. Trumpeters
forage on the ground in ocks, but roost
and nest colonially in trees. They eat both
plants and animals (including reptiles
and amphibians, and even carrion), and
sometimes follow army ants for the arthropod prey they disturb. [174]
tubenoses: Birds in the order Procellariiformes, including albatrosses, shearwaters, fulmars, petrels, and storm-petrels,
all of which have tubular nares (see separate entry). [340]
tubular nares: Nares (nostrils) that open
into a horny, tubular structure that sits on
top of the bill; in some species the tube for

Handbook of Bird Biology


each nostril is separate, but in others the

two adjacent tubes fuse to form a double
tubewithin which the tubes coming
from each nostril may remain separate
or may be partially fused. Tubular nares
are found in the Procellariiformes, an
order of pelagic seabirds that excrete
large amounts of salty uid through their
nostrils. The tube is thought to reduce
heat and airow at the nostril (where the
nasal cavity opens into the upper beak),
allowing the salty uid to ow down the
beak and away from the nostril before
evaporating. Thus the salts remaining
after evaporation are less likely to clog
the nostril and salt gland than if evaporation occurred right at the nostril. [339,
tufa: A porous, white limestone formed
under water when springs or rainwater
carry dissolved calcium into lakes that
have a high concentration of bicarbonate
ions (found in baking soda). When the
two solutions mix they form calcium carbonate, which precipitates (comes out
of solution), creating tufawhich often
accumulates into underwater towers or
crusts covering the lake bottom. Mono
Lake, in California, is well known for its
tufa formations, which became exposed
when the lakes water level dropped dramatically after water was diverted for use
in the Los Angeles area. [860]
tundra: See arctic tundra or alpine tundra. [9114]
turacos: African relatives of cuckoos, turacos are noisy and arboreal, and tend
to run squirrel-like along branches. The
plumage of turacos is soft green (one
of the few green pigments known from
birds), blue, or gray, often with bright red
(or in a few species, purple) on the wings.
The 18 species of turacos, together with
plantain-eaters and go-away-birds, make
up the family Musophagidae. [186]
turbulence: A disorderly ow of air; turbulence may interfere with smooth airow over a birds wings and thus may
disrupt ight. [513]
tutor song: A song played to a bird in the
laboratory during experiments on song
learning. [726]
tympanic canal: The lower perilymphlled canal inside the bony cochlea of
the inner ear; the tympanic canal is connected to the (upper) vestibular canal at
one end. Pressure waves in the perilymph
of the two canals are transmitted to the
endolymph of the cochlear duct between
them, setting the (in birds) basilar papilla
in motion and stimulating the sensory
hair cells. [457]
tympanic membrane: A membrane that
stretches tightly across the external ear
canal; the tympanic membrane vibrates
when struck by the pressure waves of a


sound, and transmits those vibrations to

the (in birds) columella. Also called the
eardrum. [456]
tympaniform membranes: Flexible
membranes that stretch between successive cartilaginous rings of the syrinx,
allowing the syrinx to change shape (in
response to contractions of the syringeal
muscles) and thus to produce different
types of sounds. [493]
Tyranni: One of the two large suborders of
Order Passeriformes (perching birds). The
Tyranni (also called suboscines) have less
complex voice boxes and thus sing simpler songs than members of the other passerine order, Passeri (also called oscines).
Suborder Tyranni includes birds such as
tyrant ycatchers, antbirds, woodcreepers, ovenbirds, tapaculos, manakins, and
cotingas. [725]
Tyrannidae: Extremely diverse New
World suboscine family containing
nearly 400 species; members of the family Tyrannidae are commonly known as
tyrant ycatchers. No other bird family
has more species. [180]
tyrant ycatchers: The nearly 400 species of the diverse New World suboscine
family Tyrannidae. [180]


Tym Ven

ulna: The larger and thicker of the two

bones that make up the antebrachium.
ulnare: The larger of the two wrist bones
of birds; the ulnare consists of several
fused carpals. [19]
ultimate: In biology, pertaining to longterm causes; evolutionary. Ultimate
questions about a behavior, for example, look into the factors that might
have caused the particular behavior to
evolvewhy it evolved. For comparison,
see proximate. [63]
ultimate factors: In biology, the variables
affecting the survival and reproduction
of a population and its ancestors, and
therefore inuencing the evolution of
certain traits, behaviors, or physiological
processes, or the timing of such things.
For example, the food supply at different
times of the year is a major ultimate factor that affects the timing of breeding in
birds. For comparison, see proximate
factors. [811]
ultimobranchial glands: Two small, lightcolored glands located near the parathyroid glands, usually on an artery; the
ultimobranchial glands secrete the protein calcitonin, which lowers the blood
calcium concentration. [474]
uncinate process: A attened, hookshaped extension of bone that projects
caudally from the vertebral segment of

each rib. Because each uncinate process

overlaps with the rib behind it, the processes help to strengthen the rib cage.
Found only in birds. [418]
understory vegetation: The shrubs and
shorter trees in a forest; the understory
vegetation grows below the level of the
main canopy and mid-story vegetation (see separate entry), and above the
ground vegetation. [993]
undertail coverts: The short contour feathers that cover the bases of the rectrices on
the ventral side of the tail. [16, 112]
underwing coverts: The feathers lining
the underside of the wing, from the leading edge of the wing to the ight feathers.
[112, 113]
upper beak: The upper half of the beak;
also called the upper jaw. [16]
upper jaw: See upper beak. [16]
uppertail coverts: The short contour feathers that cover the bases of the rectrices on
the dorsal side of the tail. [112, 113]
upwelling: A process by which cold, nutrient-rich water from the lower ocean
depths rises to the surface. Upwelling
may occur where certain ocean currents
meet, or where winds blowing away from
the coast move the surface water away
from shore, causing water from below to
rise to replace it. [1103, 1105]
ureters: Two tubes, one leading from
each kidney, that carry urine from the
kidneys to the (in birds) cloaca. [4125]
uric acid: A complex but less toxic molecule to which the liver of birds converts
the highly toxic nitrogenous waste products of protein metabolism. Compared
to urea, the nitrogenous waste product
of mammals, uric acid requires more energy to produce, but it is less soluble in
water and more highly concentrated, allowing birds to excrete more toxic waste
per molecule excreted, while minimizing
water loss. [4125]
urinary bladder: Sac that stores urine before it is excreted; among birds, the urinary bladder is pre-sent only in the South
American rheas. [4125]
urinary system: See excretory system.
urogenital system: The combination of
two organ systems: the urinary (excretory) system, which removes toxic nitrogenous wastes from the blood, and the
genital (reproductive) system. The two
systems often are considered together
because they are located close together
in the body, and because they may share
some structures (such as the cloaca, in
birds). [4124]
uropygial gland: See oil gland. [320]
urvogel: General term (used primarily by
the Germans) that refers to early, primi-

Cornell Laboratory of Ornithology

tive birds such as Archaeopteryx. [E4]

utriculus: One of the two membranous
chambers inside the bony vestibule of the
inner ear (the other is the sacculus); these
chambers contain hair cells and dense
crystals called statoconia, both of which
are embedded in a gelatinous material
that is surrounded by endolymph. The
statoconia and hair cells perceive the position of the head with respect to gravity
(see entry for statoconia). [458]

vagina: In birds, the short, lower section

of the oviduct, after the shell gland; the
vagina opens into the left side of the cloaca. [4130]
vagus nerve: The tenth cranial nerve; it
carries sensory and motor information
between the brain and the pharynx, larynx, heart, lungs, gizzard, liver, and intestines. [442]
vane: The broad, at surface on each side
of the shaft of a contour feather; the vanes
are formed by the interlocking of adjacent barbs. [33]
vangas: A diverse songbird family (Vangidae, 14 species) of shrike-like birds. Most
vanga species are gregarious and noisy,
gleaning insects and other small animals
as they move through the trees in groups.
vasa deferentia: See deferent ducts.
vector: 1. A quantity with both magnitude
(size) and direction; for example, a birds
ight at 25 miles per hour to the south
is considered a vector. A vector is often
shown as an arrow whose length represents magnitude and whose orientation
indicates compass direction. [568] 2.
An organism that acts as a carrier, transmitting pathogens (viruses, bacteria, and
so on) from one individual to another. For
example, certain mosquitoes are vectors
because they carry malaria from one person to another. [970]
vector navigation: Finding ones way by
adding vectors composed of compass
direction and distance in order to determine the straight-line path to a desired
destination. [579]
vein: A vessel that carries blood toward
the heart. All veins except the pulmonary
vein and its tributaries carry oxygen-poor
blood. [481, 484]
vena cava: See caval veins or cranial vena
cavae or caudal vena cava. [484]
venous ring: A ring of veins formed by the
left and right renal portal veins of birds; the
venous ring connects the various lobes of
the two kidneys, and is part of the avian
renal portal system. [483]

vent: The opening of the cloaca to the
exterior of the body. In birds, the vent is
the only posterior opening, and thus it
releases feces from the digestive system,
uric acid from the excretory system, and
sperm or eggs from the reproductive system. [4123]
ventral: Toward the belly of an organism.
ventricles: The two thick-walled, muscular, posterior chambers of the heart; the
ventricles receive blood from the atria
and then pump it to other parts of the
body: the right ventricle pumps blood
to the lungs and the left ventricle pumps
blood to the body arterial system. [477]
venules: Small veins that carry blood
from the capillaries to larger veins. [481,
vertebrae (singular, vertebra): The individual bones that form the vertebral column (backbone). [414]
vertebral canal: A long tube that runs the
length of the back, inside the backbone;
the vertebral canal is formed by the vertebral arches of successive vertebrae. The
vertebral canal protects the spinal cord,
which runs inside it from the brain to the
tail. [438]
vertebral column: Commonly called the
backbone or spine, the vertebral column
is a series of complex, uniquely articulating or rigidly fused vertebrae that provide
support for the head, neck, ribs, back,
and tail. [414]
vertebral rib: The dorsal portion of each
thoracic rib; the vertebral rib articulates
with the thoracic vertebra dorsally and
with the sternal rib ventrally. [418]
vertebrates: Animals that have a backbone: birds, sh, amphibians, reptiles,
and mammals. [11]
vertical migration: Type of migration in
which animals move seasonally up and
down mountainsides. [553]
vestibular canal: The upper perilymphlled canal inside the bony cochlea
of the inner ear; the vestibular canal is
connected to the lower perilymph-lled
canal, the tympanic canal, at one end.
Pressure waves in the perilymph of the
two canals are transmitted to the endolymph of the cochlear duct between
them, setting the (in birds) basilar papilla
in motion and stimulating the sensory
hair cells. [457]
vestibular window: A soft, oval region on
the bony cochlea, to which (in birds) the
columella of the middle ear is attached.
Movement of the columella moves the
vestibular window, sending pressure
waves through the uid (perilymph) inside the cochlea. Formerly called the oval
window. [457]
vestibule: Bony structure of the inner ear

Ven Wet
containing two membranous chambers,
the utriculus and sacculus, which contain
hair cells that perceive the position of the
head with respect to gravity. The vestibule
is part of the bony labyrinth of the inner
ear and thus contains perilymph. [448]
vestibulocochlear nerve: The eighth cranial nerve; the vestibulocochlear nerve
carries sensory input for balance and
hearing. Its vestibular (balance) portion
carries sensory input from the semicircular canals and vestibule of the inner ear,
and its cochlear (hearing) portion carries
sensations from the hair cells of the inner
ear. Formerly called the auditory, acoustic, or statoacoustic nerve. [442]
vestigial structure: A structure that has
little or no apparent function and is
thought to be present in an organism
only because it has been inherited from
an ancestor. Many vestigial structures are
reduced in size from the ancestral (and
functional) condition, and presumably
are in the process of being eliminated
through natural selection. A vestigial
structure in humans is the remains of a
nictitating membrane in the inner corner
of each eyein reptiles, birds, and some
other mammals the nictitating membrane
is functional. [346]
viduines: Refers to members of the African genus Vidua, which consists of
whydahs and indigobirds. Viduines are
colorful seed-eaters that are brood parasites on other members of their family, Estrildidae; they are known for the intricate
patterns inside the mouths of their nestlings, which strikingly resemble those of
their hosts nestlings. [8142]
villi (singular, villus): Minute ngerlike
projections. Villi often are produced by
folds in a tissue lining an organas in
the inner surface of the small intestine of
vertebrates, where the presence of villi
greatly increases the surface area available for absorbing nutrients. [4121]
visual eld: See eld of view. [451]
vitelline diverticulum: Small pouch of
tissue at the junction of the jejunum and
ileum of the small intestine in many birds;
the vitelline diverticulum is a remnant of
the yolk sac of the embryo. [4122]
vitelline membrane: In avian biology,
the transparent membrane surrounding
and holding together the yolk of an egg.
vitreous body: Clear, gelatinous material
lling the vitreous chamber (see separate entry) inside the eyeball. The vitreous body gives rigidity to the eyeball and
helps to maintain its shape. [448]
vitreous chamber: The largest chamber
inside the eyeball; the vitreous chamber
is located on the posterior side of the eyeball and is lled with a clear, gelatinous
material called the vitreous body (see

Handbook of Bird Biology


separate entry), which gives rigidity to

the eyeball. [448]
vocal repertoire: All the different types
of vocalizations that are produced by
an individual bird, including both songs
and calls; for comparison, see song repertoire. [710]
vocal signals: Vocalizations that have
evolved for a specic function. [711]
voluntary muscles: See skeletal muscles.

warm-blooded: See endothermic. [11]

warm front: The interface between a
warm air mass and the cold air mass it
is overtaking; the warm air tends to push
up over the denser cold air and become
cooler, which results in the formation
of clouds and eventually precipitation.
water birds: See aquatic birds. [165]
waterfowl: Ducks, geese, and swans;
family Anatidae. [165]
wattlebirds: Common name for members of the family Callaeidae, which
consists of two living species (the Saddleback and Kokako) and one extinct
species (the Huia). Wattlebirds are forest-dwelling songbirds endemic to New
Zealand; they are named for the pair of
colorful, eshy wattles at the corners of
their mouths. [197]
wattles: Ornamental aps or folds of
skin that dangle from the head or neck of
some birds. Wattles are found in turkeys,
pheasants, cotingas, wattlebirds, and
many others. [348]
waxbills: Together with whydahs, indigobirds, and numerous related species,
these colorful seed-eaters of the Old
World form the large family Estrildidae
(159 species). The Zebra Finch, a wellknown cage bird, is a waxbill. [183]
weavers: A diverse Old World family (Ploceidae, 117 species) of colorful
seed-eaters; weavers are found primarily in open tropical areas and are named
for the ability of many species to weave
large, complex nests. [183, 853]
well: See sap well. [9128]
western montane forest: North American coniferous forest ecosystem that is
dominated by spruce and r trees and
is found along the North Pacic slope
and in the Cascades, Sierra Nevada, and
Rocky Mountains; western montane forest hosts few breeding birds. [9116]
wetland mitigation: Compensation that
the recipient of a permit to ll or otherwise
destroy wetland areas is legally required
to provide, to make up for the wetland
area lost. Compensation may consist of



Whi Zyg

restoring or re-creating other wetland areas of equal or greater size compared to

the wetland area destroyed, or may be a
cash payment into a mitigation banka
fund managed by a public agency or
private conservation group that carries
out large-scale wetland restoration projects using money accumulated from a
large number of small-impact projects.
whisker stripe: A distinctively colored
stripe in the malar region of birds; also
called a mustache stripe or malar stripe.
[18, 210]
whistle: A song or phrase in which clear
tones are produced one pitch at a time.
whistlers: Large, round-headed songbirds with slightly hooked bills; whistlers
are known for their explosive, oftenbeautiful, songs. The nearly 40 species
of whistlers, along with shrike-thrushes,
pitohuis, and a few related species, form
the family Pachycephalidae, which is
found primarily in the Australasian region. [195]
white blood cells: Unpigmented blood
cells of several different types, all of
which contain a nucleus and can leave
the capillaries and move about in the
spaces between the body cells. Many
types of white blood cells are important
in ghting infections. Also called leukocytes. [488]
white bers: Muscle bers that appear
lighter in color than red bers because
they have fewer capillaries (and thus fewer red blood cells) and less of the reddish
oxygen-transport molecule myoglobin;
white bers also are larger in diameter
than red bers. Muscles with many white
bers are used for quick bursts of action,
but they cannot carry out sustained activity because they produce energy anaerobically (without oxygen)so lactic acid
builds up quickly and causes fatigue.
Muscles with many white bers often are
called white meat (as in the breast of a
turkey). [57]
white matter: Lighter-colored tissue
(compared to gray matter) that makes up
much of the brain and spinal cord. White
matter consists of numerous tracts, which
are bundles of nerve cell axons and their
myelin sheaths, and is found in the outer
portions of the spinal cord and the inner
areas of the brain. [439]
white meat: See white bers. [57]
whydahs: Members, along with indigobirds, of the African genus Vidua
(members of this genus are called vidu-

ines). Viduines are colorful seed-eaters

that are brood parasites on other members of their family, Estrildidae; they are
known for the intricate patterns inside
the mouths of their nestlings, which
strikingly resemble those of their hosts
nestlings. [8142]
wick: In avian biology, a beakful of vegetation that is dipped in mud and then
used to transport the mud to a nest that
is under construction; usually the entire
wick is added to the nest. [854]
wing-ashing: A foraging technique
used by some herons, egrets, and storks
in which a bird wading through the water
quickly raises or brings forward one or
both wings, apparently to frighten prey
out of hiding or to provide a shady place
where unsuspecting prey may try to hide,
thus bringing the prey within the birds
reach. [646]
wing loading: The ratio of body weight to
wing area; wing loading is a measure of
how much load each unit area of wing
must carry. [531]
wing pouch: A pocket under each wing
of male Sungrebes (Neotropical inhabitants of wooded streams); the wing pouch
is formed from a pleat of skin and is used
to carry the young while the male ies or
swims. [8137]
wing spurs: Bony outgrowths of the carpometacarpus (the main bone of the
avian manus); wing spurs are found in
various birds such as cassowaries, plovers, sheathbills, screamers, and jacanas.
winnowing: Part of a twilight territorial
and courtship display given by both male
and female Common Snipe. The birds circle high in the air and produce a series
of rapid, pulsating, whistle-hums (called
winnows) by spreading the tail and diving at high speedscausing the stiff outer
tail feathers to vibrate. European Snipe
also winnow. [717]
winter plumage: See basic plumage.
wishbone: See furcula. [419]
woodcreepers: A Neotropical family
(Dendrocolaptidae, 51 species) of similar-looking, rust-colored, suboscines
with a wide array of beak shapes; woodcreepers typically forage in tree bark
much like the unrelated Brown Creeper
of North America. [178]
woodhoopoes: A family (Phoeniculidae, 8 species) of sociable African birds
with glossy, dark plumage and long tails;
woodhoopoes nest in tree cavities and
breed cooperativelywith additional

Cornell Laboratory of Ornithology

adult birds helping the parents to tend

the nest. [185]
woodswallows: A family (Artamidae,
14 species) of small, chunky songbirds
with a graceful, swallow-like ight;
woodswallows inhabit the Australasian
and Oriental regions and are known for
huddling together on branches or in tree
cavities in groups of up to 50 or more
individuals. [196]
wood-warblers: A family (Parulidae, 115
species) of small, insectivorous birds,
many of which are colorful, found in the
New World. Many wood-warblers are
Neotropical migrants. Also called New
World warblers. [171]

yolk: In avian biology, the familiar, yellow portion of an egg; the yolk contains
nearly all of the lipids (fat) and most of
the protein needed by the developing
embryo. The yolk is surrounded and
held together by the transparent vitelline
membrane. [862]
yolking up: The deposition of yolk within the vitelline membrane in alternating
bands of darker and lighter yolk; yolking
up is part of the egg-formation process
in birds and occurs before egg laying.
yolk sac: In avian biology, the extra-embryonic membrane inside the egg that
surrounds the yolk of the developing
embryo. [865]

zooplankton: Microscopic animals and

protozoa that oat freely in aquatic environments; zoo-plankton make up the
animal portion of plankton. [1103]
Zugunruhe: See migratory restlessness.
zygodactyl feet: Foot arrangement in
which toes two and three point forward,
and the hallux and toe four point backward. Found in woodpeckers, cuckoos,
toucans, owls, Osprey, turacos, most parrots, and some other birds. [121]
zygote: The fertilized egg; the singlecelled product resulting from the union
of (1) the nucleus of the sperm cell from
the male and (2) the nucleus of the ovum
(egg cell) from the female. In birds, fertilization occurs in the infundibulum of the
oviduct. [4133

About the Authors

To ensure scientic accuracy and a high level of expertise, one or more professional ornithologists with
specic knowledge and research experience have
authored each chapter of this book. To achieve maximum clarity and readability, the twelve chapters were
carefully edited by Lab of Ornithology Education staff,
resulting in better consistency and ow among the
primary subjects. However, because each author has a
unique writing style, students may notice some differences in tone and presentation among the chapters.

Although plants rather than birds became his main interest,

his appreciation of nature derives partly from memorable experiences with birds: Once when I was a teenager, on a stormy
November day, as I walked along a hedgerow, I was astonished
to come across an isolated little tree crowded with silent birds
Id never seen beforeCedar Waxwings. Too exhausted to y,
they merely shufed down their perches a bit when I came
close. And in March of some years, the air was alive day and
night with the sound of immense ocks of Canada Geese ying
between the lake and the muddy cornelds around our house.
Experiences such as these, he notes, quicken ones appreciation
of the vivacity and fascination of the natural world, and help us
to understand what conservationists are ghting for.

Birds and Humans: A Historical Perspective Introduction: The World of Birds

Sandra G. Podulka, Marie Z. Eckhardt, and Daniel R. Otis
Sandra G. Podulka received a B.S. in Wildlife Biology from
Cornell University and an M.S. in Zoology (Animal Behavior)
from the University of Maryland, where she studied the function
of song repertoires in Song Sparrows. After graduate school,
Sandy was a Research Technician at Cornell for Dr. Stephen
T. Emlen, analyzing the social behavior of White-fronted Beeeaters, but discovered she wanted to spend more time sharing
her love of nature with others, so turned toward environmental
education. She worked at the Cayuga Nature Center for several
years, and as an Adjunct Professor of Biology at Tompkins Cortland Community College from 1988 to 1996, teaching courses
in biology and conservation. Since 1986, she has worked at
the Cornell Lab of Ornithology, in research, writing, public
education, and editingmost recently as one of the editors of
the Handbook of Bird Biology and the Home Study Course in
Bird Biology.
Sandy spent her childhood knee-deep in muddy ponds trying to catch tadpoles and frogs, and roaming elds collecting
butteriesher rst real love. She has enjoyed birds as long as
she can remember, but they did not take center stage until she
took a summer ethology course at Cornell University from Dr.
Bill Dilger. He hauled his students out before dawn every morning and taught them to recognize birds by their songs, and Sandy
has been listening to them ever since. She has participated in
Christmas Bird Counts and Breeding Bird Atlases, and has traveled to Costa Rica, Belize, Trinidad, and Peru to watch birds, but
her favorite bird-watching site is her yardwhich overlooks a
beaver pond with an ever-changing cast of avian actors.
Marie Z. Eckhardt was a biologist in the Education Program at
the Cornell Lab of Ornithology. Her academic background in
vertebrate zoology and experience as a museum consultant and
a collections manager at the New York State Museum provided
a strong foundation for her varied contributions at the Lab.
Marie has been interested in animals and the natural world for
as long as she can rememberan interest she credits to early
and regular museum visits.
Daniel R. Otis grew up on a farm in Upstate New York. Since
1988 he has worked as a freelance editor, proofreader, and writer for Living Bird and other Lab of Ornithology projects. Currently a Ph.D. student in Horticulture at Cornell, his research
focuses on conservation of the world's maple species and on
assessing the extent and effects of Norway maple invasiveness
on northeastern forests.

Dr. Kevin J. McGowan

Kevin J. McGowan is a research associate at the Cornell Lab of
Ornithology. He received his B.S. and M.S. degrees in Zoology from Ohio State University, and a Ph.D. in Biology from
the University of South Florida, where he studied the social
development of Florida Scrub-Jays. In 1988, after working as a
non-game biologist for the Florida Game and Freshwater Fish
Commission, he came to Cornell University, taking a position
as Curator of the Ornithology and Mammalogy Collections in
the Department of Ecology & Evolutionary Biology. In addition
to caring for the collections, he conducted research on crows
and taught classes in specimen preparation, eld collecting, the
relationships among birds, and Neotropical canopy biology. He
moved to his present position at the Lab in 2001.
Kevins primary research focuses on the behavioral ecology
of birds. Currently he is studying the reproductive and social behavior of two crow species in central New York State and investigating the impact of West Nile virus on crow populations. He
is an Elected Member of the American Ornithologists Union,
Webmaster and a Director of the Federation of New York State
Bird Clubs, and a member of the New York State Avian Records
Committee. Formerly, he was Secretary of the Ornithological
Societies of North America (OSNA), and editor of the Ornithological Newsletter, a bi-monthly OSNA publication.
An avocational birder since childhood, Kevin has traveled
throughout North America and to Europe, Africa, and Central
and South America to watch and study birds.

A Guide to Bird Watching

Dr. Stephen W. Kress
Stephen W. Kress is vice-president for bird conservation for
the National Audubon Society and Manager of the Societys
Maine Coast Seabird Sanctuaries. He also is ornithology Program Director for the Audubon Camp in Maine, an adjunct
faculty member in the Wildlife Department at the University
of Maine, Orono, and a Visiting Fellow at the Cornell Lab of
Ornithology, where each year he teaches the popular course
Spring Field Ornithology.
As Director of Audubons Seabird Restoration Program,
Steve advises and manages the development of techniques for
re-establishing in Maine colonies of various seabirds, such as
Atlantic Pufns; Leachs Storm-Petrels; and Arctic, Common,
and Roseate terns. In the Pacic region, he has studied the role


About the Authors

of vocalizations in attracting endangered Dark-rumped Petrels

to articial burrows in the Galpagos Islands, and Short-tailed
Albatross to decoys on Midway Island. He is author of many
books, including The National Audubon Societys Birders
Handbook, The Bird Garden, and Project Pufn, as well as
the Golden Guide to Bird Life. He also has written numerous
scientic papers on seabird biology and conservation.
During most of the year Steve lives on 33 acres of woods
and meadows near Ithaca, New York, where he manages his
land for songbirds and works on methods for restoring populations of Northern Bobwhite. He spends summers on the Maine
coast, continuing his lifelong interest in restoring nesting seabird colonies.

Form and Function: The External Bird

Dr. George A. Clark, Jr.
George A. Clark, Jr. is Professor Emeritus of Ecology and Evolutionary Biology at the University of Connecticut in Storrs.
He received his Ph.D. in Biology from Yale University, where
he specialized in Ornithology. After spending two years at the
University of Washington in Seattle, he moved to the University
of Connecticut, where he spent 32 years as a faculty member.
His 200 publications reect his research interests in the
structure, behavior, distribution, and evolution of birds. He is
past president of the Association of Field Ornithologists, served
as co-editor of the book Perspectives in Ornithology/ Essays
Presented for the Centennial of the American Ornithologists
Union, and has led educational eld trips for groups to observe
birds in North and South America, Europe, and Africa.
A birder since his high school days in Pennsylvania, George
now resides in Vermont and enjoys seeking birds by walking or
snowshoeing in the northern New England hills.

Whats Inside: Anatomy and Physiology

Dr. Howard E. Evans and Dr. J. B. Heiser
Howard E. Evans received both his B.S. and Ph.D. in Comparative Anatomy from Cornell University, where he became
a faculty member in the Veterinary College in 1950. There, he
taught gross anatomy of the horse and cow for seven years, and
anatomy of the dog, bird, and sh for 36 years. He was Secretary
of the college for 12 years and served as Chairman of Anatomy
from 1976 to 1986, when he retired. He continues to teach a
course on the literature and materials of natural history, and to
lecture in several other courses.
Howies research concerned the anatomy of reptiles and
birds, the replacement of teeth in shes, the plant-induced cyclopis in sheep, fetal development of the dog, and anatomy
of tropical shes. His most recent works include the third edition of Millers Anatomy of the Dog (1993), the fth edition of
Guide to the Dissection of the Dog (2000, with Dr. Alexander
deLahunta), and the third edition of Anatomy of the Budgerigar
and Other Birds (1996). He is co-editor of the Handbook of
Avian Anatomy, published by the Nuttall Ornithological Club
of Harvard University. He has served as President of both the
American and the World Association of Veterinary Anatomists,
and has been an associate editor of the American Journal of
Anatomy and the Journal of Morphology.
A native of New York City, Howie and his wife Erica have led
natural history trips for Cornell Adult University to the Virgin
Islands; Hawaii; Sapelo Island, Georgia; East and South Africa;
Papua, New Guinea; and Antarctica. He has lectured in China,
Russia, Taiwan, Hungary, Switzerland, Germany, England,
Thailand, Brazil, Mexico, and Japan.

J. B. Heiser is a Senior Lecturer in the Department of Ecology

and Evolutionary Biology at Cornell University. A vertebrate
evolutionary ecologist, J. B. received his B.S. from Purdue
University and his Ph. D. from Cornell University. For the past
35 years, he has taught a variety of courses in vertebrate comparative anatomy and ecology.
J. B. began teaching on the Ithaca campus, but soon was
teaching eld courses at the Shoals Marine Laboratory, an isolated island facility in the Gulf of Maine that is cooperatively
run by Cornell University and the University of New Hampshire. Eventually he became Director of the Shoals program, a
position that he held for 15 years. He won the Clark Award for
distinguished teaching and has traveled to every continent and
ocean to teach natural history on location.
Although J. B. was trained as a marine biologist and his
research focuses on the evolution and interrelationships of
coral reef sh, he has considerable natural history experience
in tropical forests worldwide. In his travels he has watched birds
(and sh) in every major biome and biogeographic region that
the planet has to offer, but he still gets a thrill out of backyard
birding in Upstate New York.

Birds on the Move: Flight and Migration

Dr. Kenneth P. Able
Kenneth P. Able is Professor Emeritus of Biology in the Department of Biological Sciences at the State University of New York
in Albany, where he had been a member of the faculty since
1971. He received his B.S. and M.S. degrees from the University
of Louisville, and his Ph.D. from the University of Georgia. Kens
research focuses on bird migration, particularly the mechanisms of orientation and navigation.
Ken has been passionately interested in birds since childhood. He has birded extensively across North America and
in Mexico, Costa Rica, Peru, Spain, France, Germany, Italy,
Switzerland, the United Kingdom, Australia, New Zealand, and
South Africa. His favorite destination is Australia, because the
avifauna is unique and the species fantastic.

Evolution of Birds and Avian Flight

Dr. Alan Feduccia
Alan Feduccia is S. K. Heninger Professor of Biology at the University of North Carolina at Chapel Hill, where he has been for
30 years. He received his Ph.D. at the University of Michigan,
where his thesis focused on the evolution of woodhewers and
Alans career has focused on vertebrate evolution, the evolution of birds, and the tempo and mode of the evolution of
modern groups of birds. His interest in the origin of birds blossomed in the late 1970s when he wrote The Age of Birds for
Harvard University Press. In 1973 he wrote a rebuttal to the
theory of hot-blooded dinosaurs in Evolution, and since that
time has been involved in the debate on bird origins. His latest
book, The Origin and Evolution of Birds (Yale University Press,
1999), addresses many of the main issues in the bird evolution
controversy, and takes what he calls the ornithological positionthat birds were already arboreal when they evolved
to y, and that they evolved from a common ancestor with
dinosaurs, but not directly from them.
Alans interest in birds began as a teenager. Later, as an
undergraduate student at Louisiana State University, he had the
good fortune to participate in bird expeditions to Honduras,
El Salvador, and Peru. He has maintained an interest in Neotropical birds ever since.

Cornell Laboratory of Ornithology

About the Authors


Understanding Bird Behavior

Dr. John Alcock
John Alcock is Regents Professor of Biology at Arizona State
University, Tempe. John is currently researching the evolution of
insect mating systems, with a special emphasis on the diversity
of male mating tactics among bee species native to the Sonoran
Desert. Earlier research has focused on birds, however, and his
textbook, Animal Behavior, An Evolutionary Approach, uses
many bird examples to illustrate all aspects of the modern study
of behavior.
John began bird watching at age ve (bird number one was
the Mallard), and seeing a good bird still boosts his heart rate. In
attempts to keep his heart rate up, he has visited Costa Rica, Ecuador, Argentina, Australia, and several countries in Europe.

Vocal Behavior
Dr. Donald E. Kroodsma
Birds enter our lives in different ways. Some of us have been
bird-crazy as long as we can remember, but others discovered
birds later. I was a late-comer, as birds grabbed me from the
chemistry lab during my last year of college. How grateful I am
that they have never let go. Immediately after college I took two
summer bird courses from the famed Olin Sewall Pettingill,
who was then director of the Cornell Lab of Ornithology but
taught at the University of Michigan eld station in Pellston,
Michigan. He put a tape recorder, headphones, and parabolic
microphone in my hands and told me to go out and tape record
some birds. In doing so, he changed my life. I went to graduate
school at Oregon State to study how young wrens learn their
songs; and then for eight years I had the good fortune to work
with Peter Marler at Rockefeller University in New York, on all
aspects of bird song. I am currently Professor of Biology at the
University of Massachusetts, Amherst. My time since college
has been spent reveling in the who, what, when, where, how,
and why of bird song.

Nests, Eggs, and Young:

Breeding Biology of Birds
Dr. David W. Winkler
David W. Winkler was born and raised in Sacramento, California. Unlike the rest of his family, he was a naturalist from
about age four. After a progression of enthusiastic interest in
butteries, wildowers, and herps, he nally settled on birds
in his early teens. He learned and studied local birds alone for
two years, when, at his rst Christmas Bird Count, he ran into
Rich Stallcup. David then spent the last two years of high school
learning a great deal from Rich about the birds of California.
While in high school, an American Birds article introduced
David to The Herring Gulls World by Niko Tinbergen, and he
carried out a senior English project on gull taxonomy within
a couple of years. By his freshman year at U. C. Davis, David
was dreaming about all the species he might someday study,
as opposed to just see, for his list.
While attending U. C. Davis, David and friends secured
National Science Foundation funding for an ecological study
of Mono Lake. This cemented his attachment to the Mono Basin and its birdlife, and his friendship with David Gaines, with
whom he co-founded the Mono Lake Committee in 1978. His
dissertation on the clutch sizes of California Gulls (with Frank
Pitelka at Berkeley) at Mono and Great Salt Lakes was followed
by post-docs at the University of Gothenburg, Sweden (with

Malte Andersson); Oxford University (with John Krebs); and

Cornell University (with Paul Sherman). David joined the Cornell faculty in 1988 upon the retirement of Tom Cade, and his
research on swallows worldwide, and Tree Swallows in Ithaca
in particular, has thrived ever since.

Individuals, Populations, and Communities: The Ecology of Birds

Dr. Stanley A. Temple
Stanley A. Temple is the Beers-Bascom Professor of Conservation in the Department of Wildlife Ecology at the University
of Wisconsin in Madison. He is also the Chair of the graduate
program in Conservation Biology and Sustainable Development in the Institute for Environmental Studies at Madison. A
quintessential Cornellian, Stan earned his B.S., M.S., and Ph.D.
all at Cornell, and for years has served as a member of the Lab
of Ornithologys Administrative Board.
Stans professional activities focus on avian ecology and bird
conservation, with a special emphasis on endangered species.
He and his students have worked with some of the worlds
most endangered birds, including Peregrine Falcons, California
Condors, Whooping Cranes, and dozens of endangered species
endemic to islands around the world. To date, none of those
species has become extinct, and most are doing signicantly
better as a result of his work.
Stan has been interested in birds as long as he can remember. Birds of prey have always been among his favorites, and
he has been a falconer for 45 years. He feels fortunate to have
incorporated all of his ornithological pleasures into his professional life.

Bird Conservation
Dr. John W. Fitzpatrick
John W. Fitzpatrick (Ph.D., Princeton University, 1978) is
Director of the Cornell Lab of Ornithology, where he arrived
in September 1995. He was Executive Director of Archbold
Biological Station, a private ecological research foundation
in central Florida, from 1988 through August 1995, and was
Curator of Birds at the Field Museum of Natural History (Chicago) from 1978 to 1989. He is a Fellow of the American Ornithologists Union (AOU). In 1985 the AOU awarded him its
highest research honor, the Brewster Award, for his book and
numerous research articles (co-authored with Glen Woolfenden) on demography, social behavior, and conservation of the
endangered Florida Scrub-Jay. Fitz has led many expeditions
to remote areas of South America, especially the western Amazonian basin and the Andean foothills. He has published numerous papers on Neotropical birds, including descriptions
of seven bird species new to science. Co-author of the book
Neotropical Birds: Ecology and Conservation (University of
Chicago Press, 1996), he has been engaged in applying science
to real-world conservation issues throughout his career. Most
recently, he helped design and implement a major network of
ecological preserves in central Florida by convening panels of
scientic experts and by engaging county, state, and federal
agencies; nongovernmental organizations; and private industry
in the process. He serves on the national governing boards of
The Nature Conservancy, the National Audubon Society, and
the Center for Biodiversity and Conservation at the American
Museum of Natural History. He is on two Endangered Species
Recovery Teams, including that of the worlds rarest bird, the
Hawaiian Crow. He enjoys watercolor painting, and has been
a bird watcher since kindergarten.

Handbook of Bird Biology


About the Authors

N. John Schmitt
N. John Schmitt is a wildlife illustrator with a lifelong interest in
birds. Since leaving the United States Army in 1973, John has
devoted his life to a variety of ornithology-related endeavors
that have taken him to many countries in Latin America, Asia,
and Europe. He has worked as a eld biologist for both the Peregrine Falcon and California Condor recovery programs.
Johns work as a eld biologist led to more serious devotion
to illustrating birds. He has illustrated several books, including the National Geographic Societys Third Edition of Birds of
North America; Clarks A Field Guide to the Raptors of Europe,
the Middle East, and North Africa; and Skutchs Birds Asleep.
Currently, he is working on eld guides to the birds of Peru
and India. John is a self-taught taxidermist/museum preparator
whose work is on display in several California museums. He
also co-leads ecotours in the United States and abroad.
A native of California, John is an avid bird watcher. His time
in the eld provides valuable inspiration and is integral to maintaining his enthusiasm. He considers his notebooks, which he
lls with written and sketch notes, as the source of his most valuable references and inspirations. John encourages everyone
interested in natural history to keep a notebook.

Cornell Laboratory of Ornithology

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