Glossary Note: In general, terms have been defined as they apply to birds. Nevertheless, many terms (especially those naming basic ana- tomical 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 defined.

Please note that this glossary is also available on the Internet at . aerodynamic valve: A vortex-like movement of air within the air A tubes of each avian lung, at the junction between the mesobron- abdominal air sacs: A pair of air sacs in the abdominal region chus and the first secondary bronchus; it prevents the backflow of birds that may have connections into the bones of the pelvis of air into the mesobronchus by forcing the incoming air along and femur; their position within the abdominal cavity may shift the mesobronchus and into the posterior air sacs. [4·102] during the day to maintain the bird’s streamlined shape during African barbets: A family (Lybiidae, 42 species) of small, color- digestion and egg laying. [4·101] ful, stocky African birds with large, sometimes serrated, beaks; abducent nerve: The sixth cranial nerve; it stimulates a muscle they dig their nest cavities in trees, earthen banks, or termite of the eyeball and two skeletal muscles that move the nictitating nests. [1·85] membrane across the eyeball. [4·41] Afrotropical region: Zoogeographic region including Mada- abiotic: Nonliving; includes both things that are dead (such as gascar, southern Arabia, and all of Africa south of the Sahara dead leaves) and those that have never been alive (for example, Desert. Sometimes called the Ethiopian Region. [1·70, 1·81] rocks). [9·7] afterfeather: A small feather that grows from the lower shaft of accessory nerve: The eleventh cranial nerve; it carries motor a contour feather and resembles the main feather but in min- output to constrict the neck muscles. [4·42] iature. [3·13] accommodation: The changes in the curvature of the lens (and age-specific fecundity: The average birth rate for females in a cornea, in birds) of the eye brought about by the action of the particular age group in a population. [9·64] ciliary muscles. These changes allow the eye to focus on objects age-specific survival rate: The proportion of individuals in a at different distances. [4·50] particular age group in a population that survive a particular acetabulum: At the hip joint, the hollow on the pelvic girdle interval of time—usually a year. [9·64] into which the head of the femur fits. [4·24] age structure: The relative proportions of individuals of dif- acoustic nerve: See vestibulocochlear nerve. [4·42] ferent ages—usually noted for a given population. [9·80] adaptation: A genetically controlled trait that increases an indi- airfoil: Any structure designed to help lift or control a flying vidual’s fitness relative to that of other individuals. [1·35] object by using the air currents through which it moves. A typi- adaptive: Describes a trait that better promotes an individual’s cal airfoil, such as the wing of a bird or airplane, is rounded on fitness than does some alternative form of that characteristic. top and curved inward below. [5·10] [6·42] air sacs: Thin-walled, transparent sacs extending from the me- adaptive management: A type of ecosystem management (see sobronchi or the lungs to different regions of the body; they act separate entry) in which managers continue to learn more about as bellows to bring air into the body and store it until expiration. the ecosystem as they proceed, and continually modify their They are found only in birds. [4·100] management techniques to incorporate the new information. air speed: A flying individual’s speed relative to the air through [10·84] which it is moving. It does not include being carried along or adaptive radiation: The evolution, from a common ancestor, slowed down by the wind, so it may or may not reflect a bird’s of a variety of different species adapted to different niches; the speed relative to the ground. [5·45] species usually have different morphologies and behaviors. albino: An individual that lacks the pigment melanin all over [1·59] its body. An individual that lacks all types of pigments is called adherent cup nest: A cup nest made of mud or saliva that re- a complete albino. [3·52] lies on chemical forces to hold it to a vertical surface; built by albumen: Egg white; albumen is composed primarily of water many swifts, including the Edible-nest Swiftlets of Southeast and protein. [8·63] Asia, whose nests are used in the Asian delicacy bird’s-nest alimentary canal: The tube for the passage, digestion, and ab- soup. [8·32] sorption of food; in most birds, it includes the esophagus, crop, adoption: In avian biology, the peaceful acquisition of a lone two-part stomach, small intestine, ceca, and the large intestine. chick or chicks by a pair of adults other than the biological It is also called the gastrointestinal tract, digestive tract, or parents. [8·128] gut. [4·103] adrenal glands: Small yellow or orange endocrine glands at the allantois: In avian biology, the extra-embryonic membrane in- cranial end of each kidney; they produce a variety of hormones side the egg that forms a sac into which the developing embryo (including adrenaline, steroids, and the sex hormones) that are shunts all metabolic wastes that cannot evaporate through the involved with circulation, digestion, and reproduction. [4·74] shell, such as uric acid crystals. [8·65, 8·69] advertising displays: Displays performed by one sex (usually Allee effect: The response, shown in some species when popu- the male) to attract a mate of the opposite sex; also called mate lation density falls below some threshold level, by which re- attraction displays. [6·37] productive behavior and/or social structure become disrupted 18 All – AOU Glossary in various ways. In some circumstances, ducts in the inner ear; it contains sensory anterior lobe of the pituitary gland: Por- this effect may cause species to be unusu- hair cells embedded in a gelatinous ma- tion of the pituitary gland that receives ally prone to extinction. [10·38] terial and surrounded by endolymph, and instructions from the nervous system in alleles: Alternate forms of genes. Most it senses changes in the animal’s speed or the form of neurohormones from the hy- animals have two alleles for each trait: direction in a particular plane of space. pothalamus. As a result, the anterior lobe one allele is on the chromosome they re- Information from all three ampullae, one secretes various hormones into the blood ceived from their mother, and the other is in each plane, is combined by the brain to that may act directly on organs or on oth- on the chromosome they received from determine the animal’s motion and thus er endocrine glands (such as the gonads, their father. As an example consider eye to aid balance. [4·58] adrenals, and thyroids); because of this color in humans, which is determined angle of attack: In a flying bird, the an- central controlling role in the endocrine primarily by whether someone has two gle between the cranial-caudal axis of system, the anterior lobe is nicknamed alleles for blue eyes, two alleles for brown the wing and the oncoming airstream. “the master gland.” [4·72] eyes, or one of each (in which case, the [5·13] anting, active: Picking up an ant or other brown dominates and the person has anisodactyl feet: Foot arrangement in chemically potent object, such as a mil- brown eyes). [10·74] which the hallux points backward and lipede, and deliberately rubbing it in the allopreening: Mutual preening during the other three toes point forward. Found feathers—presumably to deter ectopara- which two birds preen each other, usu- in most passerines. [1·21] sites. [3·22] ally around the head and neck. In many annual survival rate: See survival rate. anting, passive: Positioning oneself species allopreening not only keeps the [8·3] among a swarm of ants, permitting them plumage clean and orderly, but also to run all over the body and to move in helps to establish social bonds between ant-acacias: Various species of tropical and out among the feathers, presumably individuals. [3·19] and subtropical trees in the genus Aca- to deter ectoparasites. [3·22] cia that harbor ants inside their hollow alpine tundra: Ecosystem found above thorns. The ants receive shelter and extra antiphonal: Describes a singing interac- the tree line on mountains; it consists of nutrition from special substances pro- tion in which two individuals alternate rugged, well-drained terrain interspersed duced by the trees exclusively for the ants, their contributions. Often used to de- with meadows with low-growing vegeta- and in turn keep away insects, mammals, scribe duets between a male and female tion and a profusion of summer-bloom- and other herbivores that might feed on bird. [7·78] ing wildflowers. Very few birds breed in the trees by attacking them. The ants also antorbital fenestra: An opening on each this harsh environment. [9·114] prevent other vegetation from growing side of the skull in front of the eye socket; alternate plumage: In the Humphrey- nearby by biting off shoots as they emerge found in all archosaurs. [E·8] Parkes system of nomenclature, alternate from the ground. [8·19] antpittas: Together with antthrushes, plumage is the plumage worn by an adult Antarctic Convergence: Region of the antpittas form the suboscine family For- bird during the breeding season, if that oceans between about 50 and 60 degrees micariidae (60 species). They are small, plumage is produced by a partial molt south latitude, where cold, north-flow- drab Neotropical birds with loud, ring- before breeding. If a bird does not molt ing currents meet warmer, south-flowing ing songs; many haunt the rain forest before breeding, it continues to wear its currents, resulting in large-scale upwell- floor and may follow army ant swarms. basic plumage during breeding. In the ing of nutrient-rich water. The nutrients [1·79] traditional system, the alternate plumage support abundant plankton, which at- antthrushes: Together with antpittas, was known as the nuptial plumage or the tract a great diversity and abundance of breeding plumage. [3·33] antthrushes form the suboscine fam- seabirds. [1·104] ily Formicariidae (60 species). They are altricial: Describes young birds that antbirds: A Neotropical suboscine family small, drab Neotropical birds with loud, hatch undeveloped and in many cases (Thamnophilidae, 197 species) of small, ringing songs; many haunt the rain forest naked or with sparse down; such help- insectivorous, forest birds; some species floor and may follow army ant swarms. less young require complete parental specialize in the technique of following [1·79] care. [8·106] columns of army ants to prey on the in- aorta: The main artery exiting the heart; alula: A group of two to six feathers pro- sects and other arthropods stirred up by its branches distribute oxygenated blood jecting from the phalanx of the bird’s first the numerous moving ants. [1·79] to all parts of the body. [4·78] finger (its thumb) at the bend of the wing. It antebrachium: The middle portion of aortic arch: The curve of the aorta, just reduces turbulence by allowing fine con- the forelimb, consisting of the radius and trol of airflow over the wing. [1·11, 5·15] after it exits the left ventricle of the heart. ulna. The secondary feathers attach to the In birds, the aorta curves to the bird’s right American Ornithologists’ Union (AOU): ulna. [1·9] as it passes dorsal to the heart and toward The largest organization of professional anterior: Toward the front of an organ- the backbone, but in mammals, the aorta ornithologists in North America; it pub- ism, using the earth as a frame of refer- curves to the left. [4·83] lishes the research journal The Auk and ence. With birds, technically used only the Check-list of North American Birds AOU: See American Ornithologists’ within the eye and inner ear. But in prac- Union. [1·39] (commonly called the AOU Check-list; tice, often used interchangeably with the see separate entry). [1·39] term cranial. [1·4] AOU Check-list: Bird checklist produced amnion: In avian biology, the extra-em- by the Committee on Classification and anterior air sacs: General term referring Nomenclature of the American Orni- bryonic membrane inside the egg that be- to the air sacs nearest the bird’s front comes filled with fluid and surrounds the thologists’ Union; it contains common end—the cervical, clavicular, and ante- and scientific names of all birds that oc- developing embryo, allowing it to move rior thoracic air sacs. [4·102] and stay moist, and preventing its various cur in North America north of Mexico, or growing parts from sticking to or blocking anterior chamber: Space within the eye near North American coasts, including one another. [8·65] between the iris and the cornea; it is filled Hawaii, and is the generally accepted with aqueous fluid, which nourishes the reference for common names of North ampulla: A membranous chamber at the eye and removes wastes. [4·48] American birds. [1·64, 1·111] base of each of the three semicircular Cornell Laboratory of Ornithology Glossary Apo – Asi 19 aponeuroses: Shiny, broad sheets of con- bills and gaudy, clashing colors. They are bird and reptile characteristics, its rela- nective tissue that bind muscle fibers to- endemic to the Oriental zoogeographic tionship to modern birds and other rep- gether to form muscles. [4·27] region. [1·89] tiles has been highly controversial since aposematic: Having bright, bold colors Asities: Members (with false sunbirds) its discovery in the early 1860s. [E·2] and patterns (often reds and oranges) that of the family Philepittidae, endemic to archosaurs: All reptiles in the Archosau- advertise to potential predators that an Madagascar. These two species of subos- romorpha, the large group of diapsid individual is bad tasting or poisonous. A cine passerines feed on fruits. [1·88] reptiles that includes all thecodonts and few species that are perfectly palatable aspect ratio: The ratio of an object’s their descendants (including birds and also have evolved these warning colors length to its width. It is used to refer to crocodiles), but does not include lizards, and patterns. The strategy of sporting the shape of a bird’s wing: the larger snakes, or turtles. All archosaurs have an aposematic coloration is called apose- the aspect ratio, the more elongated the opening on each side of the skull, in front matism. [9·26] wing. [5·37] of the eye socket, called the antorbital fe- nestra. [E·8] appeasement displays: Displays given asynchronous hatching: Pattern of hatch- to decrease the aggression of another ing in which the eggs of a single clutch arctic tundra: Ecosystem found around individual; they usually consist of ste- hatch over a period of several days, re- the world in a belt extending north from reotyped postures that de-emphasize sulting in a brood of young of different the limit of trees; it consists of swampy the performer’s weapons or size and ages. This pattern occurs when incu- flatland covered with low-growing vege- expose vulnerable parts of its body. For bation begins at the time the first egg is tation, interrupted by countless shallow example, a submissive bird may point its laid. Because eggs are laid one per day, at lakes. Very few birds live in the arctic tun- beak down or away, fold its wings, lower one- to two-day intervals, the embryos of dra year round, but many migrants breed or turn away its head, point its tail down, the earliest-laid eggs have already started there, taking advantage of the long hours or adopt some combination of these pos- to develop by the time the later eggs are of daylight and numerous spring insects tures. [6·28] laid, and they hatch sooner. [6·88] for feeding their young. [9·114] appendicular skeleton: Portion of the skele- atlas: The first cervical vertebra; in birds, arctic tundra/coniferous forest ecotone: ton consisting of the sternum (breast bone), it articulates with the single occipital con- Transitional zone between the arctic tun- the pectoral girdle including the front limbs dyle on the base of the skull. (Mammals dra and coniferous forest ecosystems, (wings), and the pelvic girdle including the have two occipital condyles.) [4·16] sometimes called the northern timber- hind limbs (legs). [4·6, 4·18] line. It consists of fingers of open, stunted atria (singular, atrium): The two thin- spruce forest extending north between apteria (singular, apterium): Regions of walled, anterior chambers of the heart; bare or less-feathered skin between the fingers of low, shrubby tundra extending they receive blood returning to the heart south. [9·123] feather tracts of birds. [3·2] from the lungs (left atrium) or body (right aquatic birds: All birds with webbed atrium). [4·77] area-sensitive species: Species that re- feet that commonly swim, including the quire large tracts of habitat in order to atrioventricular valve: The valve between persist and breed successfully. [10·72] Anatidae; also, all deep-water waders each atrium and its corresponding ven- belonging to the order Ciconiiformes, tricle; it prevents the backflow of blood army ants: A subfamily of primarily such as herons and storks. Also called into the atrium as the ventricle contracts. tropical ants that are highly social and water birds. [1·65] [4·78] nomadic: colonies make daily raids in long streams of thousands of individuals aqueous fluid: A cell-free fluid, similar atrophy: To shrivel or die back; may be in structure to blood plasma; it fills the moving across the forest floor, looking for pathological or part of the normal course insects and other arthropods and flushing anterior and posterior chambers of the of development. [4·27] eye (between the lens and the cornea), out many other small creatures as they providing nourishment and waste re- attentive periods: In avian biology, peri- move. Numerous birds (see antbirds) and moval. [4·47] ods spent on the nest during incubation. some primates follow moving army ant [8·99] swarms to take advantage of the prey they arachnoid: The middle of the three vas- flush. [9·88] cularized membranes, called meninges, auditory nerve: See vestibulocochlear surrounding the brain and spinal cord. nerve. [4·42] arterioles: Small blood vessels branching The meninges provide sustenance and auditory tube: Air-filled tube leading from arteries. They carry blood from the waste removal for the cells of the brain from the middle ear to the throat; it helps arteries to the capillaries. [4·81, 4·84] and spinal cord, which are not served by to equalize the air pressure on the two artery: A vessel conducting blood away the circulatory system. [4·36] sides of the eardrum. In birds, the right from the heart. All arteries except the pul- arboreal: Living in trees. [E·18] and left tubes join and enter the roof of monary artery carry oxygen-rich blood. the mouth caudally through just one [4·81, 4·82] arboreal theory (of the origin of avian opening. In mammals, the right and left flight): Theory suggesting that flight origi- articular bone: Bone on the upper sur- tubes enter the mouth separately. Also face of each side of the lower jaw of many nated in small, arboreal (tree-dwelling), called eustachian tube. [4·57] reptile-like birds that jumped or glided vertebrates, near the caudal end; in birds, among tree branches. Proponents sug- auricular feathers: A patch of feathers it links with the quadrate bone of the up- gest that feathers first evolved to keep covering the external ear opening. Their per jaw, forming the joint between the the animals warmer in the cooler arbo- open texture protects the ear from debris jaws. [4·12] real environment, and then were used to and wind noise, yet helps to channel arytenoid cartilages: Two cartilages of extend jumps or glides. First proposed sounds into the ear. [1·8, 4·55] the larynx; they stiffen and hold the shape in 1880 by O. C. Marsh. Also called the Australasian region: One of the major of the fleshy folds surrounding the glot- trees-down theory. [E·18] zoogeographic regions of the world, tis—the opening of the larynx. [4·91] Archaeopteryx: A feathered reptile from stretching from a line termed “Wallace’s Asian barbets: A family (Megalaimidae, 150-million-year-old Jurassic limestone Line” east of the islands of Timor and Su- 26 species) of chunky birds slightly small- deposits. Because it had a mosaic of lawesi in Indonesia, southeast to New er than a Belted Kingfisher, with thick Zealand, and including New Guinea, Handbook of Bird Biology 20 Aus – Bio Glossary Australia, Hawaii, and other islands of worn by an adult bird for the longest bird and mammal species, in that indi- the mid-Pacific Ocean. Sometimes called time each year; it usually is produced viduals living in colder regions tend to be the Australian region. [1·70, 1·91] by a complete molt. In the traditional larger than those living in warmer areas. Australasian robins: A family (Eopsaltri- system, this plumage was known as the [9·16] idae, 44 species) of songbirds endemic nonbreeding or winter plumage. If a bird Bernoulli’s law: Physical law stating that, to the Australasian region. Reminiscent does not molt before breeding, it con- in any system of airflow, static and dy- of both New and Old World robins in tinues to wear its basic plumage during namic pressure must always add up to a coloration, they actually are more like breeding. [3·33] constant. Because the airflow over a mov- flycatchers, although they usually snatch basilar papilla: The lower membrane ing airfoil (such as a bird’s wing) is faster food from the ground. [1·95] of the cochlear duct in the inner ear of above than below, the dynamic pressure autonomic nervous system: A set of birds; it is coated with a layer of sensory is higher and thus the static pressure is nerves considered as a group because hair cells. Sound waves set the basilar pa- lower. Because the static pressure below of their similar functions. Acting primar- pilla into motion, causing the hair cells the wing is higher than that above, lift ily unconsciously, they innervate the to push against the tectorial membrane, is created and the bird can remain aloft. smooth muscle of the viscera, glands, triggering nerve impulses in the hair cells [5·13] and blood vessels, thus controlling the that are sent to the brain in the process of biconical: Describes an egg that is slight- automatic function of the internal organs. sound perception. In mammals, the cor- ly longer than subelliptical; also called They are under direct chemical control responding structure is called the “basilar fusiform or long subelliptical. [8·73] membrane.” [4·59] from substances circulating in the blood. bile: Substance, produced by the liver, [4·43, 4·44] BBS: See Breeding Bird Survey. [9·62] that emulsifies fats to facilitate their di- avifauna: The set of bird species living in B cells: Special white blood cells found in gestion. In birds with no gall bladder, bile a region. [1·69] the lymphatic tissues; they are important is released directly into the small intestine axial skeleton: The portion of the skel- in the immune response because they through the hepatoenteric ducts; in birds eton consisting of the skull, hyoid ap- produce antibodies. B cells are produced with a gall bladder (and mammals), bile paratus, and the vertebral column of the by the cloacal bursa and are also impor- is stored in the gall bladder and released neck, trunk, and tail. [4·6] tant in understanding the development of through the bile ducts. [4·124] AIDS in humans. [4·123] axillaries: A cluster of feathers in the bill: A bird’s upper and lower jaws, in- bird’s “armpit”; they are recognizably beak: A bird’s upper and lower jaws, in- cluding the external covering; also called longer than those lining the wing. [1·12, cluding the external covering; also called the beak. [1·6] 1·13] the bill. [1·6] bill tip organ: An aggregation of sensory axis: The second cervical vertebra. bee-eaters: An Old World family (Me- cells at the tip of both the upper and lower [4·16] ropidae, 26 species) of brightly colored beak, best developed in ducks, geese, birds with long, slender beaks.They catch sandpipers, and snipe; it is thought to axon: The cable-like, impulse-con- stinging insects in a manner similar to sense tactile stimuli during feeding. ducting, main axis of a neuron. [4·32] that of flycatchers, and then beat them to [4·57] remove the stingers before eating them. bill-wiping: A maintenance behavior in [1·85] which a bird swipes its bill sideways on B belly: In birds, part of the lower (ventral) tree branches, the ground, or other sur- babblers: A diverse family (Timaliidae, surface of the body between the breast faces, especially after eating messy foods 267 species) of gregarious, insectivo- and the vent. [1·6, 1·9] such as oily insects or suet. [3·40] rous birds, many of which have complex bend of the wing: The prominent angle binocular vision: A type of vision that social systems and breed cooperatively. at the wrist, where the bird’s wing bends produces three-dimensional images, They are found in the Afrotropical, Ori- noticeably. [1·11] in contrast to monocular vision, which ental, and Australasian regions. [1·90] benefits of philopatry hypotheses: A set produces flat images. Binocular vision back: In birds, the dorsal side of the body, of possible explanations for why cer- results when the eyes are positioned to- between the neck and the rump. [1·6, tain individual birds might forego their ward the front of the head, so that objects 1·9] own breeding in a particular breeding are detected by both eyes simultaneously. barbs: The parallel branches extending season and act as helpers at the nest of [4·51] from each side of the rachis of the feather other breeding pairs (usually their par- binomial nomenclature: The currently shaft; collectively they form the vanes. ents or other close relatives); the benefits accepted system of naming organisms, [3·3] of philopatry hypotheses focus on the devised by Linnaeus, in which each spe- barbules: Branchlets coming off both possible benefits to young adults of re- cies is designated by two words: the ge- sides of the barbs of a feather, at right an- maining with their parents. Examples of nus and the species names. [1·47] gles to the barbs and in the same plane. these hypotheses include (1) the survival bioaccumulation: The increase in the Adjacent barbules hook together, hold- of young adults may be improved when concentration of toxic or other foreign ing the vane intact. [3·3] they remain in a group, (2) by helping, substances in organisms’ bodies as a result young adults may improve the survival of taking up the substances from the envi- basal archosaurs: See thecodonts. [E·8, of close relatives (and thus increase their E·33] ronment (through plant roots, or in ingested own indirect fitness), and (3) by staying food or water) at a rate higher than that at basal metabolism: The number of calo- with their parents, young adults may in- which they are excreted from the body. ries an organism uses when completely crease their own chance of acquiring a Many toxic substances that do not occur at rest, which indicates the amount of en- superior territory, either by monitoring naturally, such as DDT and PCBs, are read- ergy needed to maintain minimal body vacancies in neighboring sites or by in- ily deposited in body tissues but excreted functions. [4·144] heriting their natal territory. [6·89] at very slow rates, because organisms have basic plumage: In the Humphrey-Parkes Bergmann’s rule: Rule describing the not evolved mechanisms to metabolize system of nomenclature, the plumage pattern of body sizes found within most them effectively. [9·126, 9·127] Cornell Laboratory of Ornithology Glossary Bio – Bro 21 bioconcentration: The stepwise increase, the eye; because no rods or cones are passerine family Ptilonorhynchidae. See found at each higher level of the food present at this spot to capture incoming bowerbirds. [6·76] chain, in the concentration of certain light, an object whose image falls on the bowerbirds: A passerine (oscine) family chemicals in the bodies of organisms. spot is not perceived. [4·49] (Ptilonorhynchidae, 20 species) of New Chemicals that usually bioconcentrate in blocking: A method of estimating the size Guinea and Australia whose polygynous the food chain are those toxins, such as of large flocks of birds by counting the males attract females by building and DDT, that tend to accumulate (see bioac- birds in a block of typical density, begin- decorating remarkably complex “bow- cumulation) in organisms because they ning at the trailing end of the flock (so ers” out of twigs and other objects. See are taken up from the environment faster that birds are not flying into the area you bower. [6·76] than they are excreted. Also called bio- are counting), and then visually superim- logical magnification. [9·125, 9·127] brachial plexus: Plexus along the spinal posing the block onto the rest of the flock cord of birds at the level of the wing; it is biodiversity: The great wealth of living to see how many times it will fit. [2·56] associated with a cervical enlargement organisms that occur on earth. [1·106] blood/brain barrier: The specialized ar- of the spinal cord. [4·39] biogeography: The study of the distri- rangement of capillaries in the brain that brachium: The upper (proximal) portion bution patterns of living things. [9·50] prevents blood from reaching the nerve of the forelimb (wing); it contains the hu- biological indicators: See indicator spe- cells, thus protecting the brain from po- merus. [1·9] cies. [10·108] tentially toxic substances that circulate in the blood. Capillaries penetrate the brain stem: See medulla oblongata. biological magnification: See biocon- meninges surrounding the brain, but do [4·37] centration. [9·126, 9·127] not reach the actual neurons of the brain. breast: In birds, part of the lower (ventral) biomass: The total mass of all the living [4·36] surface of the body, between the throat organisms in a particular population, blood plasma: See plasma. [4·86] and belly. [1·6] community, or area at a given point in Breeding Bird Survey (BBS): A count time. [9·87] blood vascular system: Alternate name for the circulatory system. [4·76] of the breeding birds of North America biome: A major terrestrial ecosystem; conducted each summer since 1967 and in North America north of Mexico, the body downs: The down feathers of adult coordinated by the U. S. Fish and Wild- nine biomes are tundra, coniferous for- birds, found under the contour feathers. life Service. Observers count birds seen est, deciduous forest, grassland, south- Body downs are most common in water or heard during three-minute periods at western oak woodland, pinyon-juniper birds and hawks, as they provide extra half-mile (0.8-km) intervals along a 25- woodland, chaparral, sagebrush, and insulation. [3·16] mile (40-km) stretch of road. Because scrub desert. Biomes are named for the bone: Tissue composed of living cells in the same routes and stops are sampled dominant vegetation of the region’s cli- a mineralized matrix; the bones provide each year, BBS data can be used to track max community, but they include all support for the body and attachment sites population trends, and the results can be the living organisms in that community, for the muscles. [4·3] correlated with habitat. [9·62] the physical surroundings, and all the bony labyrinth: The bony, outer system of breeding plumage: See alternate plum- ecological processes that occur there. fluid-filled canals (containing perilymph) age. [3·33] [9·109] that make up the inner ear. It forms the breeding season: The period of time dur- biotic: Living. [9·7] cochlea, semicircular canals, and ves- ing the year when a particular species bipedal: Walking (or running) on two tibule, and encloses the membranous may breed. [8·11] labyrinth. [4·57] legs. [E·16] bristles: Highly specialized contour bird community: See community. booming sacs: Brightly colored outpock- feathers in which the rachis is stiffened [9·82] etings of the esophagus of some North and lacks barbs along its outermost parts. American grouse that appear at the sides [3·13] birds-of-paradise: A family (Paradisaei- of the neck; they fill with air and act as dae, 46 species) of forest-dwelling song- resonators to produce loud sounds dur- broadbills: A family (Eurylaimidae, 15 birds found primarily in New Guinea; ing displays. [4·114] species) of stocky, brightly colored, sub- they are famous for the spectacular, col- oscine birds found throughout forests orful plumages and displays of the males. booted podotheca: A smooth podotheca, and scrublands of the Old World trop- [1·94] divided into long, continuous, nonover- ics, particularly in the Oriental region. lapping scales. [1·20] birth rate: The number of young born to Broadbills use their wide, flat, colorful an individual or set of individuals, or born boreal forest: Coniferous forest ecosys- bills to snatch up large insects. [1·81] into a population or species, in a given tem dominated by spruce and fir trees bronchi (singular, bronchus): The two period of time (often a year). Usually and found around the world, generally major air tubes branching off the lower (especially for humans) expressed as the in a belt north of temperate zone de- end of the trachea; one goes into each number of births per 1,000 individuals ciduous forests and south of the arctic lung. [4·92] per year. Also called fecundity. [9·49] tundra. Many birds migrate to the boreal forests to breed, taking advantage of the brooding: Sitting on hatched young, or blastoderm: In avian biology, the flattened long daylight hours and abundant insects sheltering them under the wings, primar- disc of dividing cells that lies on the upper for feeding their young. Some bird spe- ily to keep them warm, but also to protect surface of the yolk and is the first stage in cies are year-round residents. Also called them from sun, rain, or predators. Occurs the development of the embryo. [8·63] taiga. [9·116] either in the nest, or outside the nest in blind: A structure used to conceal a per- those species whose young leave the nest bounding: A flight pattern in which a bird shortly after hatching. [8·95] son so that he or she may observe birds or alternates flapping (during which it rises other wildlife; known as a “hide” in Great slightly) with glides on closed wings (dur- brood parasite: A bird that lays eggs in Britain. [2·29] ing which it descends slightly). [5·26] the nests of other species, leaving the blind spot: Site on the retina where the op- resulting young to be raised entirely by bower: A complex mating structure built the host parents. Some species parasitize tic nerve penetrates the retina and leaves by male bowerbirds—members of the Handbook of Bird Biology 22 Bro – Cen Glossary others only occasionally; others, called productivity. The tallest trees that stick cast: See pellet. [4·119, 4·120] obligate brood parasites, never build above the main canopy, somewhat like caudal: 1. Toward the tail or posterior part their own nests, and lay all their eggs in “lollipops,” are not considered part of of the body. [1·4] 2. Pertaining to the other species’ nests. [8·139] the canopy; they are called the emergent tail. [4·15] layer. [9·93] brood patch: A patch of skin on the breast caudal mesenteric vein: Vein that in birds and belly of birds that has lost feathers capillaries: The smallest blood vessels. brings blood from the lower portion of and become swollen through both the Within all body tissues except the epi- the digestive tract to the renal portal veins retention of large amounts of water in the dermis and those in the central nervous that form the venous ring connecting the tissues and the expansion of blood ves- system, capillaries form networks called lobes of the two kidneys; it is part of the sels feeding the skin. It develops a few capillary beds, which connect the ar- avian renal portal system. [4·85] days before egg laying in most individual terioles and venules. The thin walls of birds (either male or female) that incu- the capillaries allow materials to be ex- caudal vena cava: Single large vein that bate their eggs by sitting on them, and changed between the blood and the body gathers blood from veins coming from increases the efficiency of heat transfer cells—the body cells absorb oxygen from the legs, tail, kidneys, and caudal regions to the eggs. One large patch or several the blood, and the blood takes up wastes of the body and delivers the blood to the smaller ones may develop, depending on from the body cells. [4·81] right atrium of the heart. [4·83, 4·84] the species. [7·94] capillary bed: See capillaries. [4·81] caudal vertebrae: The vertebrae of the tail. In birds, the anterior caudal verte- brood reduction: Practice carried out by cardiac muscle: A special type of smooth some parent birds whose young hatch brae fuse with the sacral and lumbar ver- muscle that forms the bulk of the heart. tebrae and some of the thoracic vertebrae asynchronously and thus vary in size, Cardiac muscle can contract without in which the parents first feed the most to form the synsacrum; the next four to being stimulated by the nervous system. nine vertebrae articulate freely; and the vigorously begging offspring until it can [4·31] swallow no more, and then move on to posterior ones fuse to form the pygostyle another. Thus, in years with low food sup- carina: See keel. [4·23] (tail bone). [4·15, 4·18] plies only the largest and strongest young carinates: Birds that have a keel (carina) caval veins: General term for the right will survive, but in years with abundant on their sternum; includes all flying birds and left cranial vena cavae and the cau- food smaller young will survive as well. and many smaller flightless ones. [4·23] dal vena cava; these large veins gather Brood reduction ensures that at least carnivores: Meat-eaters. [9·123] deoxygenated blood from the body and some young will survive in years when return it to the right atrium of the heart. carotenoids: Pigments producing bright food is scarce. [8·98] [4·77, 4·84] yellow, orange, or red colors. They are Brown-headed Cowbird: See cowbird. synthesized only by plants, so birds must cavity adopters: Birds that nest in cavities [8·142, 8·148] obtain them via their diets. [3·51] but do not excavate their own, instead obtaining cavities that were created by bursa of Fabricius: Former name for the carotid artery: Main artery supplying the cloacal bursa. [4·123] physical forces (such as decay or erosion) head and neck region; among birds it is or by other species. Bluebirds, Great bustards: A family (Otididae, 25 species) highly variable, with different species Crested Flycatchers, Tree Swallows, and of large, heavy-bodied, flat-headed birds having none, one, two equal in size, or House Wrens are all cavity adopters. Also with long legs and necks. These ground- one large and one small. [4·84] called secondary cavity nesters. [8·39] dwelling birds are strong runners, and carpals: The bones of the wrist. In birds, CBC: See Christmas Bird Count. [9·60] frequent open areas in the Oriental, they are fused and reduced to just two Australasian, and southern Palearctic bones—the radiale and ulnare. [1·9] cell: Membrane-enclosed unit capable of regions, although most species live in metabolism and reproduction; the basic carpometacarpus: The largest bone of Africa. [1·83] structural and functional unit of life. [4·2] the manus of birds, formed by the fusion of some of the carpals (wrist bones) with cell body: The part of a neuron containing the metacarpals (palm bones). [1·9] the nucleus and surrounding cytoplasm; also called a soma. [4·32] C carrion: The flesh of dead animals. calamus: The hollow lower portion of the [9·28] center of origin (of a species): The geo- feather shaft, part of which lies beneath graphic area where a species evolved. carrying capacity: The maximum popu- [9·50] the skin; it has no vanes. [3·3] lation size or density that a particular area call notes: Bird sounds that are generally can support over the long term, without central fovea: Area in the central part of shorter and simpler than songs. Many any degradation in the quality of the area the retina of most birds and mammals seem to convey a specific message, such or its resources. [9·70] where the cones are most concentrated as begging calls (hunger), alarm calls and the neural layer (the nerves from the cartilage: A tissue with living cells em- rods and cones, which overlay the rods (danger), and contact calls (the caller’s bedded in a nonmineralized matrix; car- location). [7·14, 7·72] and cones and block some light) is the tilage is found in the flexible joints and is thinnest, and thus vision is the sharpest. CAM: See chorioallantoic membrane. capable of growth or resorption as well as The central foveae provide sharp mon- [8·69] transformation into bone. [4·6] ocular views of the areas to the sides of camouflage: Coloration of an organism or casques: Enlargements on the top of the bill the bird. [4·49, 4·53] structure serving to conceal it from pred- or the front of the head, usually involving central nervous system (CNS): The brain ators, other enemies, or prey. [3·61] the underlying bone. Found in cassowaries and spinal cord. [4·35] and hornbills, among others. [3·48] canopy: The upper, continuous level centrum: The main body, or central axis, of vegetation in a forest; it contains the cassowaries: A family (Casuariidae, 3 of a vertebra; the anterior end of the cen- branches of tall, mature trees and the epi- species) of large, flightless ratites inhabit- trum connects to the preceding vertebra, phytes that grow on these branches. In ing New Guinea and Australia. They have and the posterior end connects to the fol- dense forests, the canopy receives most a distinctive bony casque on top of their lowing vertebra. [4·15] of the sunlight and thus has the greatest head. [3·49]

Cornell Laboratory of Ornithology Glossary Cer – Cla 23 cere: A leathery band of skin covering checklist: A printed list of the birds found ciliary processes: Structures of the cho- the base of the bill, into which the nos- in a particular area. [2·46] roid of the eye that attach to the lens and trils open; presumably the cere protects cheek: See malar region. [1·7, 1·8] hold it in place. Ciliary muscles move the the nostrils. Present only in certain birds, ciliary processes, which in turn move the such as hawks, pigeons, and some par- chin: A small area under the lower beak lens, changing its shape. Only in birds do rots. [3·40] of birds. [1·7, 1·8] the processes attach directly to the lens. cerebellum: A large, deeply folded choana: A single slit in the roof of the [4·47] structure on the dorsal surface of the mouth, running in an anterior-posterior circadian rhythms: Daily cycles of behav- hindbrain, attached to the brain stem by direction, through which the two nasal ioral and physiological events exhibited two pairs of stout neural tracts; it con- cavities open to the mouth. [4·90] by organisms; they are regulated by an trols muscular coordination and plays an chorioallantoic membrane (CAM): internal biological clock and persist even important role in balance, posture, and Membrane covering the entire inner when organisms are kept under constant proprioception. The cerebellum of birds surface of the avian eggshell, inside the environmental conditions. Examples in- is particularly large, due to the demands inner shell membrane; it develops part- clude daily patterns of activity, body tem- of flight. [4·37] way through embryonic development perature, or nectar production. [5·61] cerebral hemispheres: The two large, from a fusion of two extra-embryonic circannual rhythms: Cycles of behavior, smooth lobes on the dorsal anterior region membranes, the chorion and allantois. growth, or other physiological activities of the forebrain; together they form the ce- The CAM is richly invested with blood that occur on approximately a yearly rebrum, which coordinates and controls capillaries, and together with the pores in basis; like circadian rhythms, they are complex behaviors, including memory the eggshell, it allows the embryo to carry regulated by an internal biological clock and learning. In mammals, these lobes out gas exchange as it receives oxygen and persist even when organisms are have folds and grooves. [4·36] from outside the egg and expels carbon kept under constant environmental con- dioxide. [8·66, 8·69] cerebrum: See cerebral hemispheres. ditions. Examples include yearly patterns [4·36] chorion: In avian biology, the extra-em- of migration, shedding and regrowth of bryonic membrane inside the egg that sur- antlers, and hibernation. [5·62] cervical air sacs: A pair of air sacs—one rounds the entire avian embryo and the circumpolar constellations: The stars sac on each side of the body—located in other three extra-embryonic membranes the neck region of birds; one sac usually near to and surrounding the North Star (the allantois, amnion, and yolk sac). It is (Polaris). [5·87] extends from each lung, but sometimes homologous (evolutionarily related) to a series of cervical sacs are located along the mammalian membrane, also called cisticolas: A family (Cisticolidae, 117 the neck, as in geese. [4·101] the chorion, which forms much of the pla- species) of small, drab, insectivorous cervical enlargement: A swelling along centa in most mammals. [8·65, 8·69] warbler-like birds of open, grassy areas of the Old World, primarily Africa. [8·7] the spinal cord at the level of the wings, choroid: The middle layer of the three associated with the brachial plexus. main layers of the eye; it lies just inside CITES: See Convention on International [4·40] the sclera. It is pigmented and forms the Trade in Endangered Species. [10·93] cervical vertebrae: The vertebrae of iris and ciliary processes. [4·48] class: Level of classification of organ- the neck region; the cervical vertebrae Christmas Bird Count (CBC): A count isms above “order” and below “phy- of birds have uniquely shaped centrum of the wintering birds of North America lum”; similar orders are placed within ends (see heterocoelous centrum ends). conducted each year since 1900 and the same class. All birds are in the class [4·15] coordinated by the National Audubon Aves. [1·52] chalazae: In avian biology, the gelati- Society. Observers count as many indi- clavicle: The collar bone; in nearly all nous, usually milky white, stringy coils vidual birds as possible within one of the birds, the left and right clavicles fuse with of albumen (egg white) that surround and circular count areas 15 miles (24 km) in a small interclavicle bone to form the V- protect the egg yolk, and are visible at ei- diameter that are scattered across North shaped furcula (wishbone). [4·19] ther end of the yolk as twisted cords. The America and beyond. Observers record clavicular air sac: A single, median air chalazae attach to the far ends of the egg- their time spent and distance covered, sac between the clavicles and surround- shell and form a suspension system for so the numbers of birds seen can be ad- ing the bifurcation of the trachea of birds. the yolk that allows it to rotate throughout justed for observer effort. The data can be [4·101] embryonic development. [8·62] used to track winter bird distribution and claw arc: The angle between the tip and channelization: The process by which abundance, as well as long-term popu- lation trends. [9·60] base of a claw, considering the claw as a humans deepen and straighten natural section (arc) of a circle; used by research- streams, converting them into water- chromosomes: Long strands of DNA ers to compare the curvatures of different filled ditches. Theoretically channeliza- found in the nuclei of most cells. Each claws. It is significant because claw cur- tion controls flooding along the stream, section of the chromosome (a specific vature is related to a bird’s habits: ground but it destroys the stream ecosystem and sequence of nucleotides) that codes for dwellers have flatter, straighter claws often increases flooding downstream. a specific protein is called a gene. [1·44, (smaller claw arcs), perching birds have [10·85] 4·134] moderately curved claws, and trunk chaparral: North American ecosystem ciliary muscles: Muscles that, in birds, at- climbers have the most highly curved found on the low hillsides of southwest- tach to the ciliary processes, which attach claws (largest claw arcs). [E·6] ern California; it consists of dense stands to the lens of the eye. When the ciliary clay lick: Clay banks rich in minerals, of broad-leaved evergreen shrubs, dom- muscles contract, they move the ciliary such as calcium, that are visited by fru- inated by chamise and manzanita. This processes, which squeeze the lens and givorous or seed-eating birds, such as ecosystem is dry, with long hot summers make it become more round. In mam- parrots and macaws, who eat the clay to with frequent fires; rain falls only in win- mals, ciliary muscle contraction relaxes obtain minerals that are otherwise lack- ter. A moderate number of birds breed in the lens, allowing it to become round by ing in their diet. Clay licks are one type the chaparral. [9·120] elastic rebound. [4·48] of mineral lick. [9·28]

Handbook of Bird Biology 24 Cle – Con Glossary Clean Water Act: Federal law passed in continental shelf, feeding mainly on fish, morphs differ in color; for example, the the United States in 1972 that attempted crustacea, and mollusks, which they find red and gray phases of Eastern Screech- to improve the quality of surface waters on or near beaches and other shorelines. Owls and Ruffed Grouse, and the blue by controlling pollution and requiring They visit land frequently, during both and white phases of Snow Geese. [9·81] sewage treatment. Section 404 (see sep- the breeding and the nonbreeding sea- columella: Small, thin bone extending arate entry) contains a set of protections son. [1·98] across the middle ear of birds, attached for wetlands. [10·93] cochlea: Elongate, bony, structure of the at one end to the inner surface of the climax community: The association of inner ear that is concerned with hearing; eardrum and at the other end to the ves- plants and animals (and other organisms) it is part of the bony labyrinth and thus is tibular window of the inner ear; it trans- that can perpetuate itself in a given area filled with perilymph. It contains the peri- mits sound waves from the eardrum to the in the absence of large-scale climatic lymph-filled vestibular and tympanic ca- fluid-filled cochlea. [4·56] change, disease, or human disturbance. nals, which are connected to one another comb: Fleshy, erect structure positioned The climax community is the final stage at one end, and the endolymph-filled co- longitudinally on top of the head of a in the process of ecological succession in chlear duct, which lies between them. In bird, often with a serrated margin (like any given area. [9·109, 9·110] mammals, but not in birds, the cochlea is a hair comb), as in domestic chickens. cline: A gradual change in certain char- coiled like a snail’s shell. [4·59] [3·48] acteristics of individuals of the same spe- cochlear duct: Membranous canal commensal: Describes a relationship cies, which is evident in a geographic inside the cochlea, bounded above by between two species or individuals in progression from one population to the the tectorial membrane and below by which one benefits and the other neither next. [1·55] a membrane that in birds is called the benefits nor is harmed. For example, cloaca: Common opening at the lower end basilar papilla. The cochlear duct is part when Cattle Egrets follow large herbivo- of the avian digestive tract for the diges- of the membranous labyrinth and thus is rous mammals to eat the insects they stir tive, excretory, and reproductive systems; filled with endolymph. [4·59] up as they move, the egrets benefit but it receives feces from the large intestine, cochlear window: A soft spot at the end the mammals appear unaffected in any uric acid from the kidneys, and eggs or of the bony cochlea farthest from the way. [9·91] sperm from the gonads, and releases these vestibular window; it acts both as a pres- communal roost: A group of birds gath- materials through the vent. [4·123] sure-release valve and as a damper for the ered to spend the night together, sleeping; cloacal bursa: A lymphoid organ that waves in the cochlea. Formerly called the may consist of just one species or a num- opens into the roof of the cloaca in young round window. [4·59] ber of different species. Birds that form birds and atrophies in later life; it pro- coevolution: An evolutionary interaction particularly large and noisy communal duces special white blood cells called between two or more species in which roosts include vultures, ravens, crows, B cells, which are important in immune one evolves an adaptation that affects starlings, herons, egrets, ibis, grackles, function and are of interest to researchers another, and then the other evolves an blackbirds, cowbirds, robins, and the for their role in the development of AIDS adaptation in response, and then the first extinct Passenger Pigeon. [6·65] in humans. Previously known as the bur- evolves another adaptation in response community: All the populations of spe- sa of Fabricius. [4·123] to the response, and so on. Coevolution cies living and interacting with one an- cloacal phallus: The copulatory organ of sometimes results in a kind of “battle” or other in the same place. Communities male ratites and waterfowl; it is an elon- “arms race” between two species. No- also can be defined as containing only gate, spiral, ridged structure that erects table examples of coevolution include certain types of species in one location, by lymphatic pressure during copulation. interactions between predators and prey, such as a forest bird community or a Sperm travel along its surface to reach the plants and pollinators, and brood para- stream insect community. [9·3] sites and hosts. [8·146] cloaca of the female. [4·129] competitive exclusion principal: Rule cloacal protuberance: A swelling at the cohort: A group of individuals born during stating that no two species can occupy caudal end of the deferent ducts, vis- the same time period (often a year). [8·5] exactly the same ecological niche in a ible externally in hand-held birds with cold-blooded: See ectothermic. [1·1] community; if they did, then eventually the feathers parted; it is present only in cold front: The interface between a mass one would outcompete the other and breeding male passerines, and is used by of cold air and the warm air mass it is over- cause it to go extinct. [9·102] researchers to determine sex and breed- taking; the dense, cold air tends to wedge complete albino: An individual lacking ing condition. The swelling is caused under the warm air, forcing the warm air all types of pigments in the plumage, by the enlargement of structures in the up and cooling it abruptly, forming pre- eyes, and skin. [3·52] terminal regions of each deferent duct cipitation that is often accompanied by complete molt: A type of molt in which during the breeding season, and may strong winds and lightning. [5·69] help to keep sperm slightly cooler than the entire feather coat is replaced. the body’s core temperature (as does the colic ceca (singular, cecum): Pouches [3·28] scrotum of mammals). [4·129] extending from the junction between conchae: Two thin, scroll-like structures the small and large intestines that hold clutch: A complete set of eggs; those laid (one median, one anterior) extending partly digested food long enough for from the lateral wall of each nasal cavity; in an uninterrupted series, for a single bacterial action to further break it down; nesting, by one female. [8·78] they are covered with a mucus-secreting digested material is released to the large membrane that contains the nerve end- clutch size: The number of eggs in a given intestine, where any released nutrients ings of the olfactory nerves, which sense clutch. [8·78] are absorbed. Birds may have none, one, odors. The mucus traps dust, and blood CNS: Central nervous system; the brain or one or two pairs of colic ceca, and vessels in the membrane warm the in- and spinal cord. [4·35] the size is highly variable among species. haled air. [4·90] Also spelled caeca or caecum. [4·123] coastal: Of the coast; coastal bird spe- cones: One of the two kinds of light-sen- cies primarily occupy the shallower wa- colies: See mousebirds. [1·85] sitive cells lining the retina of the eye; ters around oceanic islands or above the color phases: Polymorphisms in which the they are responsible for visual acuity (due

Cornell Laboratory of Ornithology Glossary Con – Cou 25 to their tight packing) and for sensing col- tection. The species are listed in three Ap- corridors: In conservation biology, long, or information (via four or five pigments pendices: Appendix I lists the most endan- narrow areas of wildlife habitat that con- and specialized oil droplets in birds), but gered species, for which all commercial nect larger areas, thus allowing individ- are not very sensitive to low light levels. trade is prohibited, Appendix II lists spe- uals to move between the larger areas. When light energy stimulates a cone cell, cies that would be in immediate danger if [10·80] it sends a nerve impulse to the brain via trade were not regulated, and Appendix III countercurrent exchange system: A sys- the optic tract. [4·48] lists species added by individual countries tem in which two fluids (liquids or gases) conical: See pyriform. [8·73] that are requesting international help in flow adjacent to one another, but in op- regulating their trade. [10·93] coniferous forest: Ecosystem of colder posite directions, while heat energy or parts of the temperate zone, where there convergent evolution: The process by materials move passively from one to the is sufficient moisture to support a forest; which organisms evolve similar forms, other, from higher to lower temperature cone-bearing trees, especially spruce behaviors, or ecological characteristics or from higher to lower concentration. and fir, dominate the vegetation. North not because they are related, but to meet An example is the countercurrent heat- America contains three main types of similar environmental challenges. [1·40, exchange system located in the legs of coniferous forests: western montane 9·106] many gulls and waterfowl: warm blood forest, boreal forest (taiga), and eastern cooperative breeding: Breeding system on its way to the feet flows through a net- montane forest. See each forest type for in which adults other than the breeding work of small vessels, which intertwines more information. [9·116] pair help the breeding pair to rear their with another network of small vessels carrying cold blood flowing from the feet coniferous forest/deciduous forest offspring. In birds, if the helpers assist with incubation or care of nestlings, they back to the body core. As a result, the ecotone: Transitional zone between the blood returning from the feet is warmed, coniferous forest and deciduous forest are called “helpers at the nest” (see sepa- rate entry). [6·88] conserving body heat. [4·151] Another ecosystems; it has a mixture of the two example is the countercurrent exchange types of trees and, in North America, has cooperative foraging: A technique in system that efficiently accomplishes more species of breeding birds than any which a group of individuals work to- gas exchange in a bird’s lungs: blood in other region. [9·123] gether to obtain prey. For example, the capillaries around the parabronchi conservation biology: An applied sci- American White Pelicans may swim in a moves in the opposite direction to the air ence that combines information gained line or semicircle and beat their wings to in the air spaces. [4·100] drive schools of fish into shallow water. through the biological fields of ecology, countershading: A type of coloration in population biology, animal behavior, Other pelicans, cormorants, and mer- gansers, among others, forage coopera- which an organism is darker on top than and genetics to attempt to reverse the below; countershading provides camou- widespread declines and extinctions of tively some of the time. Also called social foraging. [6·64] flage by reducing the contrast between species occurring throughout the world the top and shadowed underside of an today. [10·39] coracoids: Strong, stout, paired bones of organism so that it appears less three-di- conservation plan: See habitat conser- a bird’s pectoral girdle; they are not pres- mensional. [3·63] vation plan. [10·92] ent in mammals. During flight the cora- coids function as a powerful brace hold- countersinging: Interaction in which two Conservation Reserve Program (CRP): ing the shoulder joint, and thus the wing, birds sing back and forth to each other, Part of the 1985 Food Security Act that away from the body while the pectoral alternating their songs. [7·36] authorized the USDA to lease millions of muscles pull on the wing in the opposite coursers: Together with pratincoles, form areas of marginal croplands from farmers direction. [4·19] the family Glareolidae (17 species). These each year, paying them to keep the land in slender, plover-like ground nesters live in perennial vegetation to reduce soil ero- corcoracids: The two members of the pas- serine (oscine) family Corcoracidae—the open areas in tropical parts of the Old sion and crop surpluses, and to restore World, and sometimes cool their eggs or wildlife habitat. The program has helped blackbird-like White-winged Chough and the smaller, seed-eating Apostle- young by partially burying them in sand prairie songbirds by providing millions of or by bringing water to the nest in their acres of grassland habitat. [10·35] bird. These large, Australian cooperative breeders range over agricultural fields in breast feathers. [1·83] conspecifics: Members of the same spe- huge flocks when not breeding. [1·95] courting nests: Nests, usually unlined, cies. [3·66] core area: The portion of a large, continu- produced by male Winter Wrens and contact call: A sound produced by a ous habitat, such as a forest, that is far some male Marsh Wrens to attract fe- bird that appears to inform a nearby bird from the edges and thus is suitable to host males. A male may build a number of (usually a family member) of the caller’s species that would be adversely affected courting nests on his territory. In Marsh location. Often uttered by a mated male if forced to live near the edges. [9·100] Wrens, females may add a lining and lay and female as they forage relatively close eggs in the courting nest of their chosen to one another. [7·73] cornea: The transparent anterior surface mate. [8·24] of the sclera; it allows light to enter the contour feathers: Feathers that make up eye. [4·48] courtship displays: Displays performed the exterior surface of a bird, including for the opposite sex to acquire a mate of the wings and tail; they streamline and coronary arteries: Arteries (usually two) the same species, maintain a pair bond, shape the bird, and usually have well-de- arising from the first part of the aorta; they and/or stimulate and synchronize breed- veloped barbules and hooklets. [3·5] carry blood that nourishes and supplies ing behavior. [6·34] oxygen to the heart muscle itself. [4·79] Convention on International Trade in En- courtship feeding: Feeding of a female dangered Species (CITES): International coronary veins: Veins that carry deoxy- by her mate that occurs during court- agreement to which 150 nations volun- genated blood and wastes from the mus- ship and incubation, often in response to tarily subscribe that binds participating cle tissue forming the heart back into the ritualized begging calls and postures that parties to monitor, regulate, or prohibit right atrium of the heart. [4·79] resemble those of the young. The func- the import and export of species that the corpuscles: An alternate name for blood tion of courtship feeding is unknown, but group has deemed worthy of global pro- cells. [4·86] hypotheses (which are not mutually ex-

Handbook of Bird Biology 26 Cov – DDE Glossary clusive) include (1) it strengthens the pair mingos, geese, penguins, parrots, jays, cuckoo-shrikes: A diverse group of ar- bond, (2) the food improves the female’s and terns; creches of King Penguins and boreal songbirds, many of which have a condition, and (3) it allows the female to Emperor Penguins may contain several shrike-like bill and are slender and (in assess the male’s ability to provide food thousand young. Creche formation al- some cases) barred like cuckoos; how- for the young. [6·6, 8·104] lows some birds (notably penguins) to ever, they are related to neither group. coverts: The smaller feathers that partly conserve energy. It also may decrease Together with minivets and trillers, they overlie the flight feathers of the wing and predation on the young, and may free make up the family Campephagidae, tail at their bases, like evenly spaced parents to spend time foraging in distant found in the warmer parts of the Old shingles on a roof. For more information, areas, thereby allowing them to bring World. [1·96] see specific types. [1·11, 1·13] more food back for the young and to pre- cup nest: A nest in the shape of a cup, pare themselves for the next breeding. usually constructed of mud, or small cowbirds: Six species of dark, slender, [8·126] medium-sized birds that forage on seeds twigs and dried grass, with a depression and insects; they are members of the New crepuscular: Active at dawn and dusk. in the center to hold the eggs; built by the World family Icteridae (blackbirds and [9·31] majority of bird species. [8·30] New World orioles). All species except the crest: Tuft of feathers on the peak of the cursorial: Adapted for running. [E·14] Bay-winged Cowbird are obligate brood head that either stick up or can be raised. cursorial theory (of the origin of avian parasites: they lay their eggs only in the [2·10] flight): A theory suggesting that the ances- nests of other species. The Brown-headed cricoid cartilages: Two major cartilages tors of birds evolved to fly by first running Cowbird, found throughout the United that make up the sides and floor of the along the ground, and then by jumping States and much of Canada, successfully larynx. Also called the laryngeal carti- and leaping, which was augmented by parasitizes more than 140 host species and lages. [4·91] the evolution of wings and feathers, which is thought to have caused severe population eventually led to full flight. First proposed declines in some. It is the only brood par- crop: A dilation of the lower esophagus that stores food; it is found in many birds by Samuel Williston in 1879. Also called asite common in North America north of the “Ground-Up Theory.” [E·14] Mexico. [8·142, 8·148] that eat dry seeds or fruit containing seeds. [4·115] cytoplasm: The contents of a cell outside cracticids: Members of a distinctive fam- crop milk: Milk-like substance produced the nucleus but within the cell mem- ily (Cracticidae, 10 species) of songbirds brane. [1·44] endemic to Australasia. They have stout, by pigeons and doves; it is composed of straight beaks; loud, melodic calls; and a fluid-filled cells sloughed from the lining generalist, crow-like diet of small verte- of the crop and is regurgitated to feed to brates, eggs, insects, and fruits. Cracti- nestlings. Crop milk is high in lipids and vitamins A and B, and has a greater pro- D cids include butcherbirds, currawongs, dabbling: A foraging technique in which tein and fat content than human or cow and the Australasian Magpie. [1·95] a bird moves the beak rapidly on the sur- milk. Also called pigeon’s milk. [4·116] cranial: 1. Toward the head. In practice, face of shallow water to pick up small may be used interchangeably with an- cross-fostering experiments: Studies in aquatic animals and plant materials; it is terior (but see separate entry). [1·4] 2. which young of one species are placed used by “dabbling ducks” and a few other Pertaining to the head, brain, or skull. with host “parents” of another species, species. [6·47] who then rear their “adopted” young. [4·13] dabbling ducks: Ducks (such as Mallards, [9·11] cranial kinesis: The ability of the bird’s teal, wigeon, and pintails) that feed by upper jaw (upper beak) to move upward crown: The top of the head. [1·7] dabbling on the water’s surface, in con- at the same time that the lower jaw (lower CRP: See Conservation Reserve Pro- trast to diving ducks, which dive under beak) is depressed, an action permitted gram. [10·35] water to search for plant material or by the highly flexible craniofacial hinge. crus: The lower leg; in birds, supported aquatic organisms. Dabbling ducks also [4·13] by the tibiotarsus bone. [1·14] may tip “bottom up” and reach down un- der the water to obtain submerged food. cranial nerves: Twelve sets of paired cuckoo: General term for birds in the large Also called puddle ducks. [6·47] nerves, each with a specific function, family Cuculidae. The nearly 50 species serving the head, neck, and thorax region. of Old World cuckoos are in subfamily dark meat: See red fibers. [5·7] Most exit from the medulla oblongata of Cuculinae, and are all obligate brood Darwin’s Finches: See Galapagos Finch- the brain. [4·37] parasites. The familiar two-note chime es. [1·60] cranial vena cavae: Two large veins (right of the cuckoo-clock mimics the song dawn chorus: The great amount of bird and left) that gather deoxygenated blood of one of these species—the Common song heard around dawn. At this time the from large veins coming from the wings, Cuckoo of Eurasia. In the New World, largest number of bird species are sing- head, and neck and deliver it to the right cuckoos are in two subfamilies, Coccyzi- ing, and they sing more frequently, and atrium of the heart. [4·83, 4·84] nae, whose members, all nonparasitic, often more energetically and with more craniofacial hinge: Flexible joint where are termed “New World Cuckoos” and variety than at other times of the day. Why the upper beak connects to the rest of the include the Yellow-billed Cuckoo, Black- birds sing most at dawn, and in different skull. Also called the nasal-frontal hinge. billed Cuckoo, and Mangrove Cuckoo of ways than they sing during the day, is not [4·11] North America; and Neomorphinae (the known. [7·75] neomorphine cuckoos), whose members dawn song: Bird song usually given only cranium: The part of the skull enclosing are all Neotropical and include three during the early morning hours; it differs the brain; the braincase. [4·11] species that are obligate brood parasites. from a species’ normal daytime song. creche: An assemblage of the still-de- [8·141] [7·75] pendent young of two or more (usually Cuckoo-Roller: A crow-sized, arboreal DDE: A stable, persistent, toxic organic many) breeding females, attended by one bird with a stout, broad bill; it is the only compound (1,1-dichloro-2,2-bis(p- or more adults. Bird species whose young member of its family (Leptosomatidae) chlorophenyl)eth-ylene) formed in the form creches include some pelicans, fla- and is found only in Madagascar. [1·88]

Cornell Laboratory of Ornithology Glossary DDT – Dip 27 body by the metabolism of the organic ings show only in flight or when they are not lay additional (replacement) eggs if pesticide DDT (see separate entry). DDE flashed. They may have antipredator or one or more are removed from a clutch accumulates in fatty tissues and is ex- social functions. [3·64] during laying. Most determinate layers creted very slowly, and when concen- delayed plumage maturation: A situation will lay a new clutch if the entire clutch trations become high it can cause death found in some species, in which one sex is destroyed, however. For comparison, or other toxic effects such as reproductive remains in subadult plumage longer than see indeterminate layers. [8·90] failure resulting from eggshell thinning the other sex. [3·32] deterministic: In biology, describes (due to the disruption of calcium metabo- dendrites: Rootlet-like extensions from events whose occurrence and/or out- lism). Thin eggshells severely decreased comes are inevitable, based on a certain reproductive success in North American the cell body of a neuron; they usually receive nerve impulses across a synapse set of starting conditions. For compari- raptors in the 1950s and 1960s, causing son, see stochastic. [9·76] populations of most raptor species to from other nerve cells, but may transmit plummet. [10·52] them as well. [4·32] dialect: A geographic cluster of similar vocalizations (bird song, human speech, DDT: An organic pesticide (dichloro-di- dens: The upwardly projecting knob or peg of the axis (second cervical vertebra); or the sounds of other animals) that is phenyl-trichloroethane) used commonly a consequence of those vocalizations in the United States from the mid-1940s it fits into a hole in the ventral surface of the atlas (first cervical vertebra). [4·16] being learned. Dialects may exist over to the early 1970s to control Mexican very small or very large areas, depending boll weevils, gypsy moths, mosquitoes, density: See population density. [9·58] on the details of dispersal and learning. and other insect pests. DDT is highly per- density-dependent factor: A factor that [7·53] sistent in the environment and is taken in regulates population density in such a diapsid: Describes a condition in which by organisms and converted to DDE, a way that the magnitude of its effect is toxic compound that accumulates in fatty the skull has two openings on each side determined by the population density. in the temporal region, posterior to the tissues and is excreted very slowly. DDT For example, population density is de- was banned in the United States in 1972, eye socket. Diapsid skulls have this ar- creased by disease, which has a greater rangement, and diapsid reptiles are those but is still used in other countries, includ- and greater effect as population density ing Argentina, Belize, Ecuador, Guyana, reptiles that have diapsid skulls—includ- increases and thus rates of disease trans- ing thecodonts and their descendants Peru, and Mexico. See DDE for more in- mission go up. Other density-dependent formation. [9·125, 10·52, 10·53] (including birds and crocodiles), snakes, factors include predation, parasite levels, and lizards. Turtles are the only living death rate: The number of individuals in and competition for resources. [9·69] group of reptiles that are not diapsids. a species or population that die in a given density-independent factor: A factor that [E·7] period of time; also called the mortality affects population density in such a way rate. [9·49] differential exploitation: A situation in that the magnitude of its effect does not which direct interactions among two deciduous: Describing trees or shrubs depend on the population density. Ex- or more individuals that seek the same that lose their leaves during part of the amples include severe weather, natural resources are avoided because the in- year. They generally drop their leaves disasters, and the failure of food supplies. dividuals use the resources in slightly during the drier season (which in the [9·75] different ways or use slightly different temperate zone is winter) to conserve dentary: In birds, the bone forming each resources. Differential exploitation may moisture, which might otherwise be lost side of the lower beak. [4·11] result from the evolution of either mor- through evaporation. [9·117] deoxygenated blood: Blood whose red phological or behavioral differences decomposers: Level in a food chain or blood cells carry very little oxygen. Also among individuals. [9·39] web that consists of organisms (such as called oxygen-poor blood, it is found in dilution effect hypothesis: The idea that earthworms, fungi, and bacteria) that all veins except the pulmonary veins and colonial breeding may benefit individu- eat dead organisms and waste products, venules. [4·81] als because their large numbers can over- breaking them down into basic nutrients depolarizing material: A substance that whelm the consumption capacity of lo- (such as oxygen, nitrogen, and phospho- can be used to take a polarized light cal predators. [6·59] rus) and returning them to the soil for beam and vibrate it in all directions, cre- plants to use. [9·123] dinosaur theory (of bird evolution): The ating waves of all orientations, to form an theory that birds evolved from thero- decurved: Curved downward. Used to unpolarized beam. [5·94] pods such as Compsognathus approxi- describe beaks such as that of the White dermal papilla: The portion of a feather mately 150 million years ago (proposed Ibis. [1·17] papilla formed from dermal tissue; blood by Thomas Huxley in 1868) or from deferent ducts: Highly convoluted tubes vessels extend from the dermal papilla Dromaeosaurs such as Deinonychus (also called vasa deferentia) that carry into the developing feather, providing approximately 110 to 120 million years sperm from the testes to (in birds) the clo- nourishment. This core of tissue remains ago (proposed by John Ostrom in 1973). aca; in birds, the lower portion of each in a feather follicle throughout a bird’s [E·8, E·10] is enlarged to form a temporary storage life, ready to aid each round of feather de- diopter adjustment ring: The ring on bin- receptacle for sperm. [4·128] velopment after a feather is lost. [3·26] oculars, usually on one of the eyepieces definitive plumages: Any of the plumag- dermis: The inner layer of the skin; it lies but sometimes on the hinge post, that es of a fully mature bird; they may change just beneath the epidermis and contains allows the eyepieces to be focused inde- seasonally, but do not change from year blood vessels, muscles, and nerves. pendently to make up for the differences to year as the bird ages. Some species, [3·26] in visual acuity between an individual’s such as gulls, large raptors, and pelagic two eyes. [2·30] descending aorta: The continuation seabirds, take several years to reach their of the aorta (after it exits the heart and dip angle: The angle at which the mag- definitive plumage. [3·30] curves) down through the body and to- netic field lines around the earth contact deflective coloration: Conspicuous ward the tail. [4·82] the earth. The dip angle is 0 degrees at markings found on otherwise cryptically the magnetic equator, and approaches 90 determinate layers: Bird species that will colored organisms; on birds, these mark- degrees near the magnetic poles. [5·90]

Handbook of Bird Biology 28 Dir – Due Glossary direct benefits hypothesis: One pos- wing and uttering distress calls, while dorsal: Toward the back (the vertebral sible explanation for why females of slowly fluttering along the ground away column) of an organism. [1·4] some species choose the males with from the nest or young. [6·53] double-clutching: A technique in which the most elaborate ornaments (such as diurnal: Active during daylight. [9·31] biologists remove one or several eggs ornate plumage) to copulate with. The divergent evolution: A type of evolution from the nests of indeterminate layers explanation applies primarily to females and rear those eggs in the lab while the choosing nonpaternal sexual partners, in which different populations of the same species become increasingly distinct birds lay replacement eggs and rear the either (1) males for extrapair copulations, resulting young in the wild. Used by con- or (2) mates in species in which males from one another over many generations (due to exposure to different ecological servationists to increase the population do not provide parental care or other size in declining species such as Cali- resources for their offspring (such as ter- factors), eventually diverging into two or more new species. [1·57] fornia Condors and Peregrine Falcons. ritories with food or nesting sites). The [8·91] direct benefits hypothesis suggests that diving ducks: Ducks (such as Canvas- females may choose the most ornament- backs, Redheads, scaup, and goldeneyes) down feathers: Soft, fluffy feathers, typi- ed males because they are least likely to that feed by diving under the water’s sur- cally lacking a rachis. Because the bar- infect the females with mites, a disease, face to obtain aquatic plants or animals, bules lack hooks, the barbs do not cling or some other affliction. Thus the female in contrast to dabbling ducks, which re- together, so they trap more air and thus gains reproductive advantages because main at the water’s surface. [6·45] provide extra insulation. Some adult birds have body downs under their con- her health is not diminished by her mate DNA (deoxyribonucleic acid): The ge- choice. [6·84] tour feathers, and young birds have natal netic material of all cellular organisms down before molting into their juvenal direct fitness: An individual’s direct fit- and some viruses, forming the chromo- plumage. [3·16] ness is the portion of its genes that is trans- somes of these organisms. A molecule ferred into the next generation (and even- of DNA consists of two long strands of drafting: Driving closely behind another tually beyond) through the production of nucleotides, held together by bonds vehicle and being pulled forward by air its own offspring. This contrasts with in- between nitrogenous bases on the two currents moving back over the top of direct fitness, which is the portion trans- strands, with the whole structure twisted the first vehicle and swirling down and ferred as a result of an individual’s blood to form a double helix. The sequence of forward, as well as by air currents swirl- relatives producing offspring. [6·86] the nucleotides is the “genetic code,” ing up and forward from below the first which contains the instructions for mak- vehicle. Used by race car drivers or cars disjunct range: A range of a species or following trucks on a highway; the en- population that is not continuous, but ing proteins, which in turn determine the characteristics of an organism. [1·42] ergetic advantages are similar to those rather is divided into geographically experienced by a bird flying close to separate areas. [9·50] DNA-DNA hybridization: A technique and straight behind another bird, as in a dispersal: In biology, usually the move- used to determine the degree of similar- straight-line formation. [5·47] ity (in nucleotide sequence) between two ment of individuals away from the area drag: A force on a moving object, result- where they were born, or away from different samples of DNA. It is often used to compare the DNA of two different spe- ing from the friction between the object areas containing concentrations of indi- and the fluid (such as air) that it is moving viduals. [9·62] cies, to estimate how closely related they are and to hypothesize their evolutionary through. [5·16] displacement activities: Behaviors or ac- relationships. [1·46] dromornithids: A group of giant, flight- tions that seem irrelevant or inappropri- less birds that lived in Australia beginning ate to the current situation. For example, DNA fingerprinting: A technique by which the nucleotide sequences of se- in the Tertiary period, but that became a Herring Gull may stop to preen in the extinct about 10,000 years ago during the middle of a territorial conflict. Ethologists lected portions of the DNA of individu- als of the same species are analyzed and Ice Age. They are thought to be neogna- hypothesize that displacement activities thous birds most closely related to the occur because of conflicting motivations compared to determine how closely re- lated the individuals are likely to be. Used Anseriformes (ducks, geese, and swans). or indecision. Some displacement activi- [E·23] ties have become exaggerated and ritu- in criminology to determine if the DNA alized into displays. [6·32] of clues such as hair or semen match the drumming: 1. Nonvocal sounds pro- DNA of a suspected criminal; also used in duced by woodpeckers banging on dead disruptive coloration: A type of cryptic biological research to determine the relat- trees or other resonant objects with their coloration with patches, streaks, or other edness among individuals whose family beaks; most sound like short, emphatic bold patterns of color that break up the histories are not known—sometimes to drum rolls, and are given by both sexes shape of the organism, catching the eye avoid the breeding of closely related indi- to proclaim territory and attract mates. and distracting the observer from recog- viduals of endangered species. [10·76] [7·15] 2. A series of accelerating, non- nizing the whole organism. [3·62] domed nest: A cup nest with a woven vocal, muffled “thumps” at a very low distal: Away from the center of the body overhead dome that likely helps to con- frequency, produced by male Ruffed (fingers are distal to the elbow) or from ceal the eggs or nestlings; built by mead- Grouse to proclaim territory and attract the origin of the structure (the tip of a owlarks, snipe, Ovenbirds, and others. mates. Produced by repeatedly bringing feather is distal to its base—where it is [8·34] the wings forward and upward so rap- attached). [1·4] idly that with each stroke they compress dominance hierarchy: A ranking system distraction displays: Displays in which a parcel of air between the chest and of social status among members of a wings, creating a sound wave without the a bird or other animal feigns injury or in group; often established and maintained some other way creates a highly notice- wings and chest ever touching. Usually through displays and various aggressive- performed while standing on a hollow able fuss or disturbance, in order to shift a submissive behaviors, including, on oc- potential predator’s attention away from log, which acts as a resonating chamber casion, physical combat. Many domi- to amplify the sound. [7·15] the bird’s nest or young. For example, nance hierarchies are linear—A domi- Killdeer give a broken-wing distraction nates B, B dominates C, and C dominates duet: Singing performance by two indi- display by dragging and flapping one D—but other arrangements exist. [6·24] viduals. Avian duets usually are given by

Cornell Laboratory of Ornithology Glossary Dum – Edg 29 a paired male and female who may sing chians from northern New England south succession begins with bare soil or an in synchrony, overlap one another, or through Georgia and Alabama; it hosts existing community. [9·109, 9·110] alternate their songs, depending on the relatively few birds. [9·116] ecology: The study of the relationships species. Most duetting species are tropi- eclipse plumage: The set of dull-colored between organisms and their environ- cal, and include some parrots, wood- feathers worn briefly after the breeding ment. [9·1] peckers, antbirds, flycatchers, shrikes, season by some adult birds, such as ducks. and wrens. [7·78] ecoregional planning: Careful, bio- In eclipse plumage, male ducks look like logically driven planning at the level dump laying: Laying an egg or eggs in females, which do not change much in of ecoregions, with the ultimate goal of the nest of a conspecific (or sometimes, appearance. Eclipse plumage is acquired preserving the species and ecosystem a similar species). Females that dump lay by a complete molt after the breeding sea- processes (for example, fire and polli- usually also build their own nests and son, and is soon replaced through a partial nation) that occur within each ecoregion. incubate their own eggs. Wood Ducks, molt that produces the brighter colors of Ecoregional planning involves geograph- Hooded Mergansers, and some other the breeding plumage. [3·34] ically delineating ecoregions; cataloging cavity-nesting waterfowl commonly ecological constraints hypotheses: A set the ecosystems within each ecoregion; dump lay, sometimes resulting in large of hypotheses that each give an expla- identifying and mapping the most impor- numbers of eggs in one nest. Also called nation for why certain individual birds tant species, communities, and habitats; dump nesting or egg dumping. [8·76, might forego their own breeding in a par- determining potential threats to species 9·95] ticular breeding season, instead acting and sites; and then prioritizing species, duodenum: The U-shaped first loop of as helpers at the nest of other breeding communities, and the key habitats as to the small intestine, running from the pairs (usually their parents or other close which require the most urgent conser- stomach to the jejunum. [4·122] relatives). The ecological constraints hy- vation action. Ecoregional planners then dura: The outermost, toughest, and potheses focus on the possible costs to set conservation goals, outline plans, and most fibrous vascularized membrane young birds of dispersing from their natal monitor the results. [10·86] surrounding the brain and spinal cord. territory; examples of these hypotheses ecoregions: Regions of the world that are Along with the other meninges, it pro- include (1) few vacant territories of good biologically distinct in terms of climate vides sustenance and waste removal for quality may be available, (2) few suitable conditions, topography, soil types, and the cells of the brain and spinal cord, breeding partners may be available, and plant communities; also called physio- which are not served by the circulatory (3) the birds may have little chance of graphic regions. [10·86, 10·87] system. [4·36] reproducing successfully until they gain “parenting” experience. [6·89] ecosystem: Both the living and nonliving dust-bathing: Driving fine particles components of a particular area (includ- through the feathers by rolling the body, ecological isolating mechanisms: Struc- ing the physical surroundings), as well as fluffing the feathers, and wiping the head tural, physiological, or behavioral adap- the ecological processes that bind them and bill in a dusty area or by picking up tations that have evolved in species that all together (such as decomposition, soil dust and throwing it over the body, af- have very similar niches, and that allow erosion, and water and nutrient cycling). ter which dust is shaken or preened out; the species to divide up the resources in Ecosystems may be small (for example, important for feather maintenance and various ways (such as using slightly dif- the ecosystem of a rotting log) or large (a removal of ectoparasites. [3·21] ferent resources or foraging in different deciduous forest ecosystem). [9·109] parts of the habitat), and thus to coexist. dynamic pressure: The pressure of a For example, certain wood-warbler spe- ecosystem management: Understand- flowing fluid, or of movement through cies can coexist in spruce forests of north- ing and maintaining entire ecosystems a fluid such as air or water. You feel dy- eastern North America even though they (see separate entry), instead of focusing namic pressure when wind blows against all eat similar small insects, because they on particular species or habitat types. your face. [5·12] forage in different parts of trees. [9·103] [10·83] dynamic soaring: A type of flight in which ecological niche: See niche. [9·102] ecotone: A zone of transition between birds use the gradient in wind speed that two ecosystems, such as the oak savanna exists over the surface of the ocean to ecological release: An expansion of the ecotone between the eastern deciduous travel for long distances without spend- niche of certain populations of a species, forests and prairie grasslands of North ing much of their own energy: the bird such that a greater breadth of resources America. Ecotones host a greater num- glides down the gradient at an angle, then such as habitat and food are used, in ar- ber of species than either adjacent eco- turns and abruptly rises into the wind, us- eas where interspecific competition is system, because some species in each of ing its momentum to gain height quickly, lower, as on islands. (In these situations, the two adjacent ecosystems frequent the then turns and glides down again, cross- the niche is “released.”) [9·104] transition zone, and because some spe- ing the ocean in large zig-zags. Dynamic ecological succession: The process by cies prefer the greater variety of resources soaring is used most by albatrosses and which one association of plants and ani- found at habitat edges and thus live spe- other large pelagic birds with high-as- mals is replaced by another, then that one cifically in those areas. [9·93] pect-ratio wings. [5·44] is replaced by another, and so on until ectoparasites: Parasites, such as flies, the climax community for that area is ticks, fleas, lice, and mites, inhabiting the reached. The types of communities, and exterior of a host’s body. [3·19, 3·23] the order in which each succeeds the pre- E vious one, is fairly predictable for a given ectothermic: Describes organisms that eardrum: See tympanic membrane. habitat and geographical region. Primary must rely on sources of heat outside their [4·56] succession begins on a substrate, such as own bodies to keep warm; also called cold-blooded. [1·1] eastern montane forest: North American rock, sand, or lava, that has never before coniferous forest ecosystem dominated supported a community. The process by edge effect: The tendency for areas near by spruce and fir trees and found below which a lake gradually fills in to form the edge of a habitat patch to differ from alpine tundra but on the higher sum- a bog community also is considered a areas near the center in a number of dif- mits, ridges, and slopes of the Appala- form of primary succession. Secondary ferent ways. For example, areas where two habitats meet fairly abruptly (such as Handbook of Bird Biology 30 Egg – ESA Glossary a forest/field boundary) often host a high- it is the only member of its family (Dromi- a bird’s shoulders, such as the red feathers er number of species than do either of ceidae) and is the second largest living on a Red-winged Blackbird. [3·5, 6·73] the two adjacent habitats. Other ways in bird, next to the Ostrich. [1·93] EPC: See extrapair copulations. [6·79] which the edge of a habitat patch may dif- enantiornithine birds: See opposite epidermal collar: Structure formed dur- fer from the center include changes in mi- birds. [E·20] croclimate, such as amount of sunlight or ing feather development as a feather pa- humidity; increased habitat disturbances, Endangered Species Act (ESA): Federal pilla elongates: the outer cells harden, such as fire and wind damage; and higher law passed in the United States in 1973 fuse, and form a ring or “collar” of epi- numbers of introduced plants or animals that commits the government to take ac- dermal tissue surrounding the dermal (because these species generally invade tion to prevent the extinction of native portion of the original papilla. The collar from the edge). Forest birds breeding near species and to protect their habitat. It cells multiply to produce most structures edges may experience higher rates of pre- also establishes a procedure to develop in the developing feather. [3·26] dation and/or brood parasitism than birds a list of threatened and endangered spe- epidermis: The outer layer of the skin; it breeding near the center, because preda- cies, identify their critical habitat, and protects the inner layer—the dermis— tors such as raccoons, squirrels, and Blue develop and carry out Recovery Plans. and does not contain blood vessels. In Jays and parasites such as Brown-headed [10·23, 10·91] birds, it gives rise to the feathers and to Cowbirds are more common near edges. endemic: Found only in a particular re- the horny sheath covering the bill, legs, [9·93, 10·73] gion; describes a species or other taxo- and feet, including the claws. [3·26] egg: 1. The ovum; the female reproductive nomic group. For example, kiwis are epiphytes: Plants that grow on other cell sometimes called the egg cell, both endemic to New Zealand. [1·70] plants but, in contrast to parasitic plants, before and just after it is fertilized by a endocrine glands: Structures that secrete use their roots for attachment rather than sperm cell. [4·130] 2. The hard-shelled hormones directly into the blood; the to obtain nutrients from their support structure laid by birds, containing the hormones are carried to other parts of the plant; examples include many bromeli- embryo, yolk, and white. [4·130] body, where they stimulate or regulate ads (“air plants”) and orchids, and a few egg tooth: A short, pointed, calcareous the activities of other glands or organs. cacti. [9·88] structure on the tip of the upper beak (and The major endocrine glands of birds are epizootic: 1. A disease that spreads sometimes the lower beak as well) that the pituitary, thyroids, parathyroids, ul- quickly among crowded animals, such as develops in bird embryos shortly before timobranchials, adrenals, gonads, pan- viral enteritis (duck plague), which may hatching; the embryo rubs and pounds creas, pineal, thymus, and cloacal bursa. kill hundreds or thousands of waterfowl. the egg tooth against the inner wall of [4·71] [9·69] 2. Term used to describe a dis- the eggshell to break it open and hatch. endocrine system: Organ system that ease that spreads quickly among animals The egg tooth sloughs off or is resorbed acts with the nervous system to initiate, when they are highly crowded. [9·69] by the growing chick within a few days coordinate, and regulate body func- epoch: A unit of geological time. Succes- after hatching. [8·104] tions, including reproduction and de- sive epochs make up a period. [1·113] velopment. It consists of the endocrine electrophoresis: A method of separating equilibrium theory: Theory proposing large, charged molecules of different glands and their secretions, called hor- mones. [4·69] that the number of species on an island lengths or charges (DNA fragments or at any one time represents a balance be- proteins treated to carry a charge) by their endolymph: The fluid that fills the struc- tween the number of new species colo- rate of movement through a thin slab of tures forming the membranous inner lab- nizing the island (immigration) and the gel in an electric field. [1·43] yrinth of the inner ear: the cochlear duct, number of species becoming extinct. A elephantbirds: A family (Aepyornithidae) the semicircular ducts, and the utriculus consequence of this relationship is that of huge, flightless ratites that lived on the and sacculus. [4·57] smaller islands tend to have fewer spe- island of Madagascar beginning 10 to 20 endothermic: Having the ability to cies because (1) they tend to have lower million years ago, but were exterminated generate one’s own body heat through immigration rates (they are less likely to about 2,000 years ago by human activity. metabolic processes; also called warm- be “discovered” by colonists), (2) they The tallest stood about 10 feet (3 m) and blooded. Only birds and mammals are have higher extinction rates (the popu- weighed about half a ton. [E·23] endothermic. [1·1, 9·13] lations are smaller and thus more likely to elliptical wings: Short, broad wings hav- entoglossal: The bone (part of the hyoid go extinct due to stochastic factors, and ing a low aspect ratio; they allow great apparatus) that supports the tongue; also the fewer resources and reduced habitat maneuverability, but do not promote called the tongue bone. [4·14] diversity are more likely to lead to com- efficient or rapid flight. Elliptical wings petitive exclusion), and (3) they typically environment: The surroundings of an contain a lower diversity of habitat types are common in birds that live in forests, organism, including physical features, woodlands, or shrubby areas, such as than larger islands, thereby providing chemical and energetic factors, and oth- niches for fewer kinds of species. This crows, grouse, quail, and most song- er living organisms. [9·7] birds. [5·37] theory applies only in situations in which environmental stochasticity: The ten- speciation is not a major source of new emarginate tail: Tail shape in which the dency of nearly all environments to ex- species, for example over short periods rectrices become slightly longer from the perience many random (or at least unpre- of time, or on islands too small or too far inside out. Also called notched. [1·19] dictable) events, such as natural disasters from other islands to permit speciation embryo: A developing young animal that or severe weather. If a population is small through geographic isolation. [9·95] is still inside its egg or mother; in some and “unlucky” enough to be struck by era: A unit of geological time. Eras are animals, especially mammals, refers one of these events, it may be entirely divided into periods. [1·113] only to the earlier developmental stages. wiped out. [9·77] [8·64] erythrocytes: See red blood cells. enzyme: A protein that catalyzes (assists) [4·88] emigration: The movement of individuals a biochemical reaction without being out of a population. [9·49] consumed in the reaction. [1·41] ESA: See Endangered Species Act. [10·23, 10·91] Emu: A large, flightless ratite of Australia; epaulettes: Patches of colored feathers on Cornell Laboratory of Ornithology Glossary Eso – Fea 31 esophagus: Thin, straight muscular tube but in no other birds, a urinary bladder protrude the tongue, lower the lower carrying food from the pharynx to the is also present); it removes toxic nitroge- beak, constrict the neck, and (in birds) stomach. It is lined with mucus-secreting nous wastes from the blood by produc- tense the columella ear bone. It also may glands that moisten food to ease its pas- ing, storing, and excreting urine. In birds, carry some taste sensory input from the sage, but contains no digestive glands. the urine is composed of uric acid. Also tongue. [4·41] [4·113] called the urinary system; it is often con- facultative partial migration: See partial ethology: The study of animal behavior, sidered together with the reproductive migration. [5·56] system as the urogenital system. [4·125] primarily from a proximate approach, fairy-bluebirds: Two species of arboreal by comparing similar behaviors in re- exit pupil: The space through which songbirds endemic to the Oriental zoo- lated species to understand how certain the light beam exiting the eyepiece of geographic region; they are named for behaviors evolved, and by investigating binoculars or a telescope passes; this the brilliant blue and black plumage of releasers and instinctive behaviors and is, in effect, the hole through which the the males, and feed primarily on figs and the underlying physiological processes. observer looks. The diameter of the exit other fruits. Together with leafbirds, they Some of the earliest animal behaviorists pupil is calculated by dividing the size of make up the family Irenidae. [1·89] (primarily Europeans) were concerned the objective lens by the magnification; mainly with ethology, and the two pio- the larger the exit pupil, the brighter the fairywrens: An Australasian family (Me- neers of the field, Konrad Lorenz and image. [2·35] luridae, 26 species) of cooperatively Niko Tinbergen, laid the groundwork breeding, wrenlike birds with long, extensor: A muscle that pulls one bone cocked tails. [1·95] for animal behavior studies carried out away from another bone. [4·27] today. [6·7] false sunbirds: Members (with asities) external ear: The portion of the ear con- of the family Philepittidae, endemic to eustachian tube: See auditory tube. taining the external ear canal and the ear- [4·57] Madagascar. These two species of sub- drum (tympanic membrane. [4·56] oscine passerines feed on insects and eutrophication: The changes in a lake, external ear canal: The channel through nectar. [1·88] estuary, or slow-moving stream when which sounds enter the ear; it leads from family: Level of classification of organ- it receives excess plant nutrients, espe- outside the body to the eardrum. [4·56] cially nitrates and phosphates. Some nu- isms above “genus” and below “order”; trient input occurs naturally, through the extinction events: Large-scale extinc- similar genera are placed in the same erosion of soil and run-off from adjacent tions of many species; they have oc- family. The scientific names of bird fam- land. In many cases, however, human curred periodically throughout Earth’s ilies end in “idae” (for example, Corvi- activity greatly increases the rate of nu- history. [1·113] dae). [1·52] trient input or adds new sources, such as extra-embryonic membranes: Mem- FAP: See fixed action pattern. [6·8] discharges from industries and sewage branes that protect and nourish the grow- fascia: Connective tissue binding to- treatment plants; when human activity ing embryo, but do not become part of gether hundreds or thousands of muscle is involved, the changes are sometimes the adult body. The four main ones in the fibers to form a skeletal muscle; it may be termed cultural eutrophication. One im- avian egg are the yolk sac, amnion, al- in the form of bandlike tendons or broad, portant change is the dramatic increase lantois, and chorion. [8·65] shiny sheets called aponeuroses. [4·27] in the growth of plants and (especially) extrapair copulations (EPCs): Copu- fat bodies: In birds, yellowish fat depos- algae, which can choke out native plants. lations with birds other than one’s mate. its laid down just under the skin, usually Then, as the masses of plants and algae [6·79] in individuals storing fat in preparation die and sink to the bottom they decom- eyeball: The eye; in birds it is flat to tubu- for migration; the most conspicuous fat pose, depleting the water of oxygen, and lar, in contrast to the spherical eyeball of bodies lie over the abdomen and in the thus few aquatic organisms can continue mammals. Its outer layer is the sclera, and depression formed anterior to the breast to survive. In addition, some algae may the entire eyeball sits in the socket and is muscles where the clavicles fuse to form produce toxins, dyes, or odors that de- protected on its exposed side by the eye- the wishbone, and are visible in a hand- crease water quality. [10·56] lids and nictitating membrane. [4·47] held bird with the feathers parted. Re- eventual variety: Pattern of singing in which eyebrow stripe: A distinctively colored searchers often use the degree of fat ac- a bird repeats one song type many times be- line running from the upper beak toward cumulation in a migrant as an indication fore switching to a different type, which it the back of the head, located ventral to of its energetic condition and potential to then repeats many times, and then switches the boundary of the forehead and crown; continue migration. [5·64] to another type, and so on. [7·86] also called the superciliary line. [1·7, feather comb: See pectinate claw. evolution: A change over time. The evo- 2·10] [3·46] lution of living things is the set of cumu- eyeline: A distinctively colored line that feather papillae: Small bumps covering lative changes in the characteristics of a passes through the eye. [2·10] the surface of the skin of birds during species or population over successive embryonic development. They consist generations that result from natural se- eye ring: A circle of distinctively colored of a core of dermis and a covering of epi- lection acting on the genetic variation feathers or skin surrounding the eye. dermis, and each will eventually form an among individuals. [1·34] [2·10] embryonic feather. Also simply called excavators: In avian biology, bird species papillae. [3·26] that dig their own nest cavities or tun- feather sheath: A thin, cylindrical tube nels, either in sandy soil (such as Belted of keratin surrounding and protecting a Kingfishers, Bank Swallows, bee-eaters, F facial disc: Flat, relatively round, forward- developing feather. It eventually breaks mot-mots, and Crab Plovers) or in wood facing part of the head of owls; it probably open to let the mature feather unfurl. (woodpeckers). [8·39] funnels sounds into the bird’s ear open- [3·26] excretory system: Organ system con- ings. Also spelled facial disk. [4·55] feather tracts: Areas of a bird’s skin sisting of the kidneys and their ducts, the facial nerve: The seventh cranial nerve; where feathers are attached; also called ureters (in the rheas of South America, it carries motor signals to muscles that pterylae. [3·2] Handbook of Bird Biology 32 Fec – Fur Glossary fecal sac: A tough, flexible bag enclosing fledging: Term commonly used to de- food web: See food chain. [9·123] the feces of most passerine nestlings; it scribe the time at which nestlings that foot: In birds, refers to the portion of the allows the parents to remove and dispose are reared in the nest leave the nest, even leg distal to the tibiotarsus bone, and of the feces more easily—parents some- though their flight abilities may not yet has two sections. The upper section is times grab the fecal sacs as they emerge be well developed. But, the term is some- supported by the tarsometatarsus bone, from a nestling’s cloaca. Many parents times used to describe the time at which a which does not touch the ground when carry the sacs some distance from the young bird has finished acquiring its first the bird walks; the lower section consists nest and drop them, but others consume complete set of flight feathers—generally of the phalanges of the toes, upon which them. [8·136] the time at which it is capable of flight. the bird walks. [1·14] fecundity: See birth rate. [9·49] The term is used less often in precocial species that leave the nest shortly after foramen triosseum: An opening at a bird’s female-defense polygyny: Mating system hatching, but sometimes it refers to the shoulder joint, formed by the junction in which males compete fiercely for con- time at which they begin to fly. “Fledg- of the scapula, coracoid, and clavicle trol of clusters of nesting females. In some ing” may also be used to refer to the pro- bones. This hole acts as part of a pulley species, such as Montezuma Oropendo- cess of reaching the moment of fledging. system that allows the force of the supra- las of Central America, a male dominance [8·115] coracoideus muscle to be redirected: hierarchy results with the top few males because the tendon of the supracoracoi- securing most of the matings. Because fledging period: The period of time from deus passes through this hole, the muscle males do not help to rear the young, this hatching to the moment of fledging (see can be located ventrally yet still raise the system may evolve when male parental separate entry). [8·116] wing. Also called the triosseal canal or care is less important to the survival of the fledgling: A young bird that has recently supracoracoid foramen. [4·20, 5·6] young than are safe nesting sites or rich fledged (see separate entry for fledging). forebrain: The anterior portion of the food supplies. [6·74] [8·115] brain; it consists of the two cerebral femur: Bone that supports the upper hind flexor: A muscle that pulls one bone to- hemispheres with the olfactory lobes at limb (thigh) of many vertebrates, includ- ward another bone. [4·27] their anterior ends. [4·37] ing birds and humans. [1·14] flight feathers: The remiges of the wings forehead: The front of the head, from the fibula: The thinner of the two lower hind and rectrices of the tail. [1·11, 1·12, crown to the base of the bill. [1·7] limb bones in many vertebrates, includ- 1·13] forked tail: Tail shape in which the rec- ing humans; in birds, the fibula is reduced flight songs: Songs given by birds dur- trices become abruptly longer from the and present only as a thin, needlelike ing flight; they are particularly common inside out. [1·19] bone running two-thirds of the way down among birds of open areas, such as grass- the side of the tibiotarsus. [1·14, 4·25] formation: An ordered arrangement of a lands and the tundra, where few perches group of birds in flight, such as V-shaped field of view: 1. The width of the area are available. Singing from higher up gen- flocks of geese or single lines of Brown visible (usually at 1,000 yards from the erally increases the distance over which Pelicans and cormorants. [5·47] observer) through binoculars or a tele- a song can be heard. Nevertheless, some scope. If the field of view is labeled in species that do have ample perches, such frequency: Also known as pitch, the degrees, multiply degrees by 52.5 to get as the forest-dwelling Ovenbird, also give frequency is the rate at which a sound the width in feet. 2. The view attained flight songs; these usually begin with a causes the air through which it is moving by a particular species, due to the place- jumble of notes that appear to draw atten- to compress and thin (one compressing ment of its eyes; also called an organism’s tion to the singer, and then proceed with and thinning is called one cycle), and is visual field. [2·37, 4·51] the bird’s normal song. Their function is measured in cycles per second, or Hertz unknown. [7·84] (Hz). The more cycles per second, the filoplumes: Hairlike but relatively stiff higher the frequency, and the higher the feathers having a rachis but few or no floaters: Animals, generally males, that pitch. [7·4] barbs (any barbs are present only at the do not hold territories or form pair bonds, tip). In the skin next to their follicles they but cruise around areas containing terri- fright molt: See shock molt. [3·38] have sensory receptors that allow them torial individuals, waiting for a chance to frogmouths: A family (Podargidae, 14 spe- to monitor movement within the feather take over a territory or sneak a copulation cies) of nocturnal forest birds of the Orien- coat. [3·17] with a paired bird. [9·70] tal and Australasian regions that resemble fitness: An individual’s degree of success flowerpeckers: A family (Dicaeidae, 43 their smaller nightjar relatives in both cryp- at contributing its genes to the next gen- species) of small, busy, noisy songbirds, tic appearance and behavior. [1·91] eration; often measured as the number primarily of the Oriental zoogeographic frontal plane: A plane, usually horizontal, of its offspring that survive to reproduce. region, that forage high in trees on ber- through an organism, dividing the body [1·35, 9·7] ries, nectar, and insects. [1·90] into dorsal and ventral portions. [1·4] fixed action pattern (FAP): A behavior follicle: A small, epidermis-lined pit in frugivorous: Feeding mainly or exclu- that occurs in complete form each time the skin of a bird, from which a feather sively on fruits. [1·81] the animal encounters the releasing emerges and to which it is attached. [3·2, functional response: A change in the stimulus—even upon the animal’s first 3·26] amount of a certain type of prey taken exposure to that stimulus. An FAP may food chain: The sequence in which or- by a predator, as a result of a change in be a simple or complex behavior, or a ganisms in an ecosystem feed upon other the population density of that prey. For series of behaviors, but once begun, it is organisms. Most food chains consist of example, as insect populations in an played out to the end regardless of any producers, consumers, and decom- area increase during spring, American response that occurs or any intervening posers. A food chain is a fairly linear and Kestrels may begin to eat more insects stimuli. [6·8] simplistic model of feeding relationships and fewer voles. [9·25] flanges: See oral flanges. [3·43, 8·107] in communities, which are more realist- furcula: V-shaped bone of the pectoral flank: The side of a bird, dorsal and cau- ically represented as food webs because girdle of birds, formed by the fusion of the dal to the leg. [1·6] of the numerous interconnections among right and left clavicles with a small inter- species and levels. [9·123] Cornell Laboratory of Ornithology Glossary Fus – Goo 33 clavicle bone. The furcula is also called is able to use a wide range of some type grabs it from the surface. In hover glean- the wishbone. [4·19] of resource; for example, animals with ing, practiced by kinglets, phoebes, and fusiform: Describes an egg that is slightly generalist diets eat many different types Great Crested Flycatchers, among others, longer than subelliptical; also called bi- of foods. [9·5] the bird hovers while taking food from the conical or long subelliptical. [8·73] genetic bottleneck: The loss of genetic di- surface of vegetation. [6·44] versity experienced by most populations glenoid fossa: In birds, a cup-shaped as they become very small. Such loss oc- depression formed where the coracoid curs for three main reasons: (1) some al- and scapula meet; it receives the rounded G leles are lost simply by chance, when the end of the humerus, forming a ball-and- Galapagos Finches: A group of 15 finch only individuals that possess them die, (2) socket joint that enables the humerus to species in the family Emberizidae living the rate at which new alleles arise in the rotate freely around the shoulder joint. in and near the Galapagos Islands. They population declines, because there are [4·20] are a classic example of adaptive radia- fewer individuals in which new mutations glial cell: See neuroglia. [4·36] tion, as they have a wide array of beak can occur, and (3) inbreeding occurs more sizes and shapes and are all thought to frequently because fewer genetically dis- gliding: Unpowered flight (no thrust is have evolved from a common ancestor. similar individuals are available as poten- provided) in which the flying object loses Also called Darwin’s Finches. [1·60] tial reproductive partners. [10·75] altitude. In a bird or other animal, flying without flapping the wings or limbs, while gall bladder: Small organ for storing bile; genetic structure: The relative propor- losing altitude (as compared to soaring, it is located under the liver and is not tions of individuals with different genetic in which the animal rises). [5·36] present in all birds. [4·124] types—usually noted for a given popula- globular nest: A spherical dome nest gallinaceous birds: Grouse, quails, tur- tion. [9·80] with a top that completely encloses the keys, pheasants, and all other birds in genital system: The reproductive system. nest; usually entered through a hole on the order Galliformes; includes domestic [4·127] the side. Examples include the nests of chickens. [1·66] genus (plural, genera): Level of classifica- Cactus Wrens, Black-billed Magpies, gametes: The ova (egg cells) and sperm tion of organisms above “species” and and Southern Penduline-Tits. [8·36] cells. [4·134] below “family.” Genus is always capi- glossopharyngeal nerve: The ninth cra- ganglia (singular, ganglion): Aggregations talized, and is underlined or printed in nial nerve; it carries sensory and motor of nerve cell bodies; they form nerve cen- italics. [1·47] information between the brain and the ters outside the central nervous system geographic range: The geographic area tongue, pharynx, esophagus, and throat. (in the peripheral nervous system). [4·36, within which a species or population It also carries motor output to the salivary 4·44] generally remains at a particular time of glands. [4·42] gaping: In avian biology, begging behav- year; a species may have different breed- glottis: Small, slit-like opening to the ior of young birds that begins shortly after ing and nonbreeding ranges. Also called larynx; it is surrounded by fleshy folds hatching in which they open the mouth the range. [2·19, 9·49] whose muscles regulate the passage of air widely; may be accompanied by a beg- germinal spot: In avian biology, the light- into the respiratory system. [4·91] ging call. Given by altricial young and colored site on the egg yolk where the glycogen body: A gelatinous mass of those precocial young whose parents embryo will eventually develop. The ger- neuroglial cells rich in the nutritive sugar feed them. [8·107] minal spot sits atop a cylinder of light- glycogen; it is located in the rhomboid gas exchange: The movement of gases be- colored yolk that stretches from the yolk’s sinus, and its function is unknown. The tween an organism and the environment; core to its surface. [8·63] rhomboid sinus and glycogen body are for example, in the lungs of many organ- gizzard: The lower part of the bird’s two- unique to birds. [4·39] isms including birds, the blood takes up part stomach; it is rounded and has a gonads: The primary sexual organs, the oxygen from the air and discharges car- tough lining and thick, muscular walls, testes and ovary, which produce, re- bon dioxide and water. [4·89] often with internal ridges. It grinds and spectively, sperm and eggs. They also are gastric cuticle: Leathery or sandpaper-like softens foods, and in birds that eat seeds, endocrine glands, secreting the sex hor- material that forms the lining of the giz- the gizzard has more muscular walls than mones testosterone, estrogen, and, from zard; it is a combination of carbohydrate in birds that eat meat. Seed-eating birds the ovary only, proges-terone. [4·75] and protein secreted by glands in the wall may eat grit or small stones, which re- Gondwanaland: The southern land mass of the gizzard. Also called koilin. [4·119] side in the gizzard to aid in grinding. In birds that eat fish or other meat, the giz- formed 200 million years ago when Pan- gastrointestinal tract: See alimentary gea split into two large land masses. It canal. [4·103] zard molds indigestible material, such as bones and feathers, into compact balls consists of present-day South America, gene: The sequence of base pairs within (pellets) that are then ejected through the Africa, Madagascar, India, Australia, a molecule of DNA that codes for one mouth. [4·118] New Zealand, and Antarctica. [1·68] specific protein. [1·41, 4·134] gleaning: A foraging technique in which good genes hypothesis: One possible ex- gene flow: The movement of genetic ma- a bird takes insects and other small inver- planation for why females of some species terial between populations. In mobile tebrates from the surface of vegetation choose the males with the most elaborate animals, gene flow generally occurs as or other substrates. In perch gleaning, ornaments (such as ornate plumage) to individuals emigrate, immigrate, or breed practiced by many wood-warblers and copulate with. The explanation applies with individuals from other populations. In other species, a bird grabs prey without primarily to females choosing nonpa- organisms such as plants and fungi, gene flying from its perch. In sally gleaning, ternal sexual partners, either (1) males flow occurs as spores, pollen, or seeds are practiced by birds such as Red-eyed for extrapair copulations, or (2) mates in carried by water, wind, or animals. [1·53] Vireos, chickadees, titmice, and some species in which males do not provide gene pool: All the genes existing in a small flycatchers, the bird sits still and parental care or other resources for their population at a given time. [6·22] watches the surrounding vegetation until offspring (such as territories with food or nesting sites). The good genes hypothesis generalist: In biology, an organism that it sees an insect move, then flies out and Handbook of Bird Biology 34 Gra – Haw Glossary suggests that the most ornamented males [6·28] in one or a few specific types of habitats have genes that increase their own sur- guano: The accumulation of seabird (habitat specialists). [10·59] vival in some way (for example, they may droppings, particularly at a breeding habitat imprinting: The process by which have greater skill at foraging, avoiding colony, dried to a hard, crusty state. a young animal, especially a bird, learns predators, or obtaining good territories), Guano is often mined and used as fertil- the characteristics of appropriate habi- and that females choose them because izer. [1·104] tat by observing its surroundings while it then their own male and female offspring is still living within its parents’ territory. may inherit those traits and have an in- guild: In avian biology, a group of eco- logically similar (but not necessarily re- When adult birds eventually settle on creased chance of surviving and produc- their own territories, most choose to live ing offspring of their own. [6·84] lated) birds that use the same resources in similar ways. For example, Henslow’s and/or breed in areas similar to those in graduated tail: Tail shape in which the Sparrows, Grasshopper Sparrows, East- which they were raised. [9·11] rectrices become abruptly longer from ern Meadowlarks, and Bobolinks all nest habitat richness theory: The idea that the outside in. [1·19] on the ground in extensive grasslands, smaller islands generally hold fewer spe- granivorous: Seed-eating. [9·27] and thus are considered to be in the same cies than larger islands because they tend gravity: The attractive force between two “grassland nesting guild.” [9·102] to have fewer different habitats, and thus masses of matter; this force, for example, guineafowl: Gregarious, chicken-like species immigrating to smaller islands are tends to draw objects toward the center birds with distinctively spotted and less likely to find suitable habitats than of the earth. [5·10] striped plumage; they are often domesti- those colonizing larger islands, and are more likely to perish (or leave). [9·96] gray matter: Darker-colored tissue cated or found in zoos. The six species are (compared to white matter) that makes endemic to Africa and form the subfam- habitat specialists: Species that live and up much of the brain and spinal cord. It ily Numidinae of the family Phasianidae. breed successfully in only one or a few consists of numerous nuclei, which are [1·85] specific types of habitats. If all else is collections of nerve cell bodies, and is gular fluttering: Opening the bill wide equal, these species are more likely to found at the core of the spinal cord and in and vibrating the thin, expansive gular go extinct than species that can live and the outer areas of the brain. [4·39] membranes of the throat, in order to dis- breed successfully in a wide range of habitats (habitat generalists). [10·59] grazing: A foraging technique in which sipate heat. This cooling method is used an animal bites off clumps of grass or oth- by pelicans, cormorants, herons, owls, habitat specificity: How wide a range of er vegetation; used by geese, antelope, and nighthawks. [4·153] different habitat types in which a given and others. [6·46] gular region: Upper part of the throat, just species can live and breed successfully. [10·62] greater coverts: The feathers partly over- below the chin. [1·7, 1·8] lying each remex on the upper surface of gut: See alimentary canal. [4·103] habituation: The permanent loss of a re- the wing. A greater primary covert over- sponse as a result of repeated stimulation lies each primary feather, and a greater without reward or punishment; learning secondary covert overlies each second- to ignore unimportant stimuli. [6·9] ary feather. [1·12, 1·13] H hacking: The technique of introducing habitat: The physical surroundings in gross primary productivity: See primary young, captive-raised birds of prey, es- which an organism lives. It consists of productivity. [9·87] pecially falcons, to appropriate habitat physical factors, such as light, tempera- by releasing them from an enclosure that ground-rollers: A family (Brachypteraci- ture, and moisture, as well as living organ- serves as an artificial nest and in which idae, 5 species) of solitary, terrestrial in- isms, such as plants and animals. Habitats biologists continue to place food until sect-eaters with stout bills, short wings, are often characterized by a dominant the bird has learned to hunt on its own. and moderately long legs and tails. They plant type or physical feature, such as a [10·100] are endemic to Madagascar and are de- grassland habitat or stream habitat. [9·8] clining in number dramatically. [1·88] hallux: The first toe, composed of two habitat conservation plan (HCP): A plan phalanges. In nearly all birds, it points ground speed: The speed of a moving ob- that must be submitted to the U. S. Fish backward. The hallux is well developed ject (such as a flying bird) in relation to and Wildlife Service by anyone who ap- in perching birds and is reduced or ab- the ground; it is equal to the bird’s speed plies for a permit to destroy endangered sent in many running birds. [4·25, 4·26] with respect to the air (the air speed) plus species or their habitats (as allowed under Hamerkop: A stork-like water bird whose or minus the wind speed, depending on a 1982 amendment to the Endangered shaggy, crested nape and stout, tapering the direction the bird is flying with re- Species Act). The plan must specify the bill make the head appear hammer- spect to the wind. [5·68] steps that the applicant will take to min- shaped. It builds an enormous mound imize the number of individuals killed ground-up theory (of the origin of avian nest. It is endemic to Africa, and is the and to minimize the impact on the spe- flight): See cursorial theory. [E·14] only member of its family, Scopidae. cies as a whole, and also must explain group selection: Theory proposed by V. [1·85] why other alternatives are not feasible. C. Wynne-Edwards in 1962 suggesting hand: See manus. [1·9] that different groups of individuals might [10·92] experience differential survival based on habitat fragmentation: The process by hatching: Emerging from the egg. A variations among the groups, and that this which a large, continuous habitat is clutch may hatch synchronously (all at might be one way in which natural selec- broken into a number of small, isolat- about the same time—see synchronous tion worked—passing down through the ed patches by activities such as devel- hatching) or asynchronously (over a pe- generations a greater proportion of the opment, logging, or farming. [9·97] riod of several days—see asynchronous hatching). [6·88] genes of the groups with higher survival habitat generalists: Species that can live rates. Theorists have basically disproved and breed successfully in a wide range hatchling: A newly hatched animal. the idea as originally presented, but are of different habitats. If all else is equal, [8·61] still exploring whether some other form these species are less likely to go extinct Hawaiian Honeycreepers: A group of 32 of group selection might be plausible. than species that can live and breed only species, many of them extinct, in the sub- Cornell Laboratory of Ornithology Glossary Haw – Hon 35 family Drepanidinae, within the family the liver to the small intestine. [4·121] [2·30] Fringillidae. These small, colorful birds herbivores: Organisms that eat primarily Hoatzin: An odd-looking, leaf-eating have a wide array of beak shapes and are plants. [9·123] bird that nests in branches overhanging a dramatic example of adaptive radiation, Hertz (Hz): A measure of the frequency lakes and slow-moving streams in the as they all are thought to have evolved Neotropics; it is the only member of its from a common ancestor. [1·62] of a sound, in cycles (of compression and thinning of the air) per second. [7·4] order and family (Opisthocomidae). To hawking: A foraging technique in which avoid predators, nestlings may tempo- a bird sits very still on a high or exposed heterocoelous centrum ends: The saddle- rarily leave the nest and clamber about perch, and when it sees an insect, flies shaped, interlocking ends of the centrum in trees, aided by small claws on their out and snatches it in midair, returning (main body) of avian cervical vertebrae. wings. [1·77] Because the anterior end is concave in to the same or a nearby perch; used by Holarctic: A combination of the Nearctic many flycatchers, kingbirds, bee-eaters, a lateral direction and the posterior end is concave in a dorso-ventral direction, and Palearctic zoogeographic regions; waxwings (sometimes), and some wood- the Holarctic encompasses the Northern peckers, among others. [6·44] articulating vertebrae can rotate freely against one another, allowing the neck Hemisphere north of the tropics. [1·70] HCP: See habitat conservation plan. to be highly flexible. [4·15] homeostatic mechanisms: Mechanisms [10·92] heterodactyl foot: Foot arrangement in that act in various ways to preserve bal- heading: The compass direction in which which the third and fourth toes point for- ance. For example, various physiologi- a bird is pointing its beak and propelling ward and the hallux and second toe point cal factors keep the body temperatures of itself through the air. Because of cross- backward; found in trogons. [1·21] birds and mammals relatively constant. winds, the heading may not be the actual And, various ecological factors keep direction that the bird is progressing with heterogametic: Describes individuals many populations at a density that fluc- respect to the ground. Also applies to oth- whose two sex chromosomes are differ- tuates around a fairly stable level. [9·67] er flying animals and aircraft. [5·68] ent; in birds, the chromosomes are called ZW and heterogametic individuals are homeothermic: Describes animals that heel pad: Calloused enlargement of the females, but in mammals, the chromo- are able to keep their internal body tem- upper end of the tarsus (at the heel), found somes are called XY and heterogametic perature constant even when the outside in the nestlings of many cavity nesters, individuals are males. Heterogametic temperature varies. In birds and mam- such as woodpeckers and trogons. It is individuals are capable of producing mals, the hypothalamus monitors body thought to reduce abrasion of the tarsus two different types of gametes (eggs or temperature and, if required, triggers re- from the rough lining of the nest cavity. sperm)—one with one type of sex chro- sponses that warm or cool the individual [3·44] mosome and one with the other. For com- to bring it back to normal body tempera- helpers at the nest: Adult birds that are parison, see homogametic. [4·136] ture. [9·13] not currently breeding themselves, but as- heterozygous: Possessing two alleles that homing ability: The ability to return to sist other breeding pairs (usually, but not are different for a given gene; for example, a specific place. For example, homing always, their relatives—especially their having one allele for blue eyes and one pigeons can return to their lofts when re- parents) in rearing their offspring. Some allele for brown eyes. (Each allele comes leased from great distances away. [5·80] helpers are birds whose own breeding from a different parent.) For comparison, homogametic: Describes individu- attempts have failed, whereas others are see homozygous. [10·74] als whose two sex chromosomes are unpaired birds or those without territo- high-aspect-ratio wings: Wings that are the same; in birds, the chromosomes ries; helpers often breed on their own in are called ZZ and homogametic indi- subsequent years. Bird species in which long, narrow, and unslotted; the length derives primarily from the lengthened viduals are males, but in mammals, the helpers are common are called coopera- chromosomes are called XX and the tive breeders. [6·88] inner wing (as compared to high-speed wings, in which the length results pri- homogametic individuals are females. hepatic portal system: Pattern of circula- marily from the long outer wing). High- Homogametic individuals are capable tion in which blood from the capillary aspect-ratio wings are highly efficient of producing only one type of gamete beds of the small intestine is brought by at producing lift at relatively high flight (eggs or sperm)—one that contains the the hepatic portal vein to the capillary speeds, but they are difficult to ma- same type of sex chromosome as the par- beds of the liver, for further processing. neuver, especially during take-offs. They ent. For comparison, see heterogametic. [4·85] are found in a few seabirds that are highly [4·136] hepatic portal vein: Large vein carrying specialized for dynamic soaring over the homozygous: Possessing two identical blood from the upper part of the small ocean, such as albatrosses, shearwaters, alleles for a given gene; for example, hav- intestine, where the blood has absorbed petrels, and some gulls. [5·42] ing two alleles for blue eyes. For compari- digested nutrients, to the liver, where it high-speed wings: Wings that are tapered, son, see heterozygous. [10·74] undergoes further chemical processing pointed, and in many cases sweptback, honeyeaters: A huge, diverse Austral- before returning to the heart. “Portal” with unslotted primaries and a high as- asian family (Meliphagidae, 181 species) veins carry blood between two capillary pect ratio; found in birds such as falcons, of dull-colored, arboreal songbirds with beds—in this case, the capillary beds in swifts, swallows, terns, ducks, and many medium-length, curved bills. Honeyeat- the small intestine and the liver—instead shorebirds. High-speed wings allow ers have a distinctive brush-tipped tongue of between a capillary bed and the heart. good control and high speeds, but are for gathering nectar and are important [4·85] energetically expensive to use because pollinators, but they also eat insects and hepatic veins: Two large veins (left and the bird must flap constantly. [5·38] fruits. They feed busily, and often congre- right) that carry blood from the capillary hindbrain: The posterior portion of the gate at flowering trees. [1·95] beds of the liver to the caudal vena cava brain; it includes the cerebellum and me- honeyguides: A nonpasserine family (In- near its entry into the right atrium of the dulla oblongata (brain stem). [4·36] dicatoridae, 17 species) of the warmer heart. [4·85] hinge post: Central rod of binoculars, parts of the Old World, whose members hepatoenteric ducts: In birds that lack a around which the two barrels pivot. are peculiar in their ability to digest wax, gall bladder, ducts that transport bile from especially beeswax, in addition to their Handbook of Bird Biology 36 Hoo – Inc Glossary insect prey. Some honeyguides lead hu- tween two individuals of different spe- immediate variety: Pattern of singing in mans or other mammals to bee nests, and cies. [3·66] which a bird sings a song type once, then then eat the wax that is exposed as the hydrology: The study of water: its distri- moves on to a different song type and mammal breaks open the nest. [1·87] bution, properties, and patterns of flow sings it once, then goes on to yet another hooklets: Tiny hooks found on each of the on the earth and in the atmosphere. type, and so on, such that the bird sings barbules that branch from the distal side [10·85] many different songs in its repertoire be- fore ever repeating one. [7·86] of each barb of a contour feather; hook- hyoid apparatus: A V-shaped unit com- lets catch onto the barbules branching posed of bones and cartilage, located immigration: The movement of indi- from the proximal side of the next barb between the two halves of the lower viduals into a population. [9·49] (toward the feather tip), lightly holding jaw. The hyoid apparatus consists of the imperforate septum: A condition in which the barbs together to form a smooth, con- tongue bone (entoglossal) and the horns the nasal septum (tissue separating the left tinuous vane. [3·4] of the hyoid, which, respectively, support and right nasal cavities) has no opening. hormone: Chemical substance secreted the tongue and the muscles that control In contrast with a perforate septum (see into the blood and thus carried to other tongue movement. [4·13] separate entry), an imperforate septum ap- parts of the body, where it may stimulate hyperphagia: The dramatic increase in pears to decrease an animal’s sensitivity in or regulate the activities of glands or or- the amount of food that birds consume detecting odors. However, it increases the gans. Hormones are the messengers of as they prepare to migrate; because these ability to locate an odor’s source, because the endocrine system. [4·69] birds eat more than their body requires in odors entering one nostril do not mix with hornbills: A family (Bucerotidae, 55 spe- the short term, they store body fat to be those from the other. [4·90] cies) of large, toucan-like birds of the Ori- metabolized for energy during migration. Important Bird Areas (IBA) Program: A ental and Afrotropical regions; hornbills [5·63, 9·24] program to identify sites (called IBAs) in have long tails and huge, down-curved hyperthermia: Condition in which the each state that are particularly important bills topped by a distinctive casque. They body temperature rises a few degrees to local populations of breeding, winter- are famous for the female’s nesting be- above normal; if the animal cannot bring ing, or migrating birds. The program gath- havior—she seals herself inside the nest its temperature down, it soon dies. [9·18] ers input from bird watchers and profes- cavity with the eggs and young. [1·91] sional ornithologists, designates the sites hypoglossal nerve: The twelfth cranial it feels are most important, and then en- horns of the hyoid: Part of the hyoid ap- nerve; it combines with the vagus (tenth) paratus; the two bones on each side of courages public and private stewardship and glossopharyngeal (eleventh) cranial of the sites. Developed as an international the hyoid that extend backward (cau- nerves to form a combined trunk that dally) from the tongue bone, running be- effort by BirdLife International, the pro- controls movement of the tongue, larynx, gram is conducted in North America by neath the skull and then curving upward trachea, and syrinx. [4·42] around the back of the head. [4·13] the National Audubon Society. [10·113] hypothalamus: The ventral portion of the imprinting: A type of early learning in House Finch eye disease: See myco- brain region between the forebrain and plasmal conjunctivitis. [9·71] which a young animal quickly acquires midbrain. The hypothalamus plays a ma- specific information for certain experi- hover gleaning: See gleaning. [6·44] jor role in hormonal control of body pro- ences. For example, after hatching, Grey- hovering: A type of flight in which an cesses by using special neurosecretory lag Goose goslings rapidly learn to follow aircraft or flapping animal remains sus- neurons to control the pituitary gland, the first moving object they encounter, pended in air in one place. In birds, hov- which extends from a stalk below the which is usually their parent. [6·12] hypothalamus. [4·36, 4·72] ering usually is achieved by positioning inattentive periods: In avian biology, the body nearly vertically and beating the hypothermia: Condition in which the blocks of time spent off the nest during wings more or less horizontally, produc- body temperature drops below normal. incubation. [8·99] ing just enough forward thrust to balance If the animal cannot bring its temperature wind speed, and just enough lift to com- back to near normal, it soon dies. [9·16] inbreeding: The mating of closely re- lated individuals within a population. pensate for gravity. Hovering is an ener- hypothesis: A tentative explanation for getically expensive undertaking. [5·26] The greater the degree of inbreeding, the an observation or phenomenon; hy- greater the chance that deleterious ge- humeral patagium: A flap of skin extend- potheses are usually stated in a way such netic traits will occur in the population. ing from the brachium to the trunk of the that they can be tested via the scientific See inbreeding depression. [9·66] bird. [1·11] method. [6·5] inbreeding depression: The reduction in humerus: The bone supporting the bra- Hz: See Hertz. [7·4] the average fitness of offspring born to chium (upper arm or wing). [1·10] parents that are closely related to each Humphrey-Parkes Nomenclature: The other, compared to the fitness of offspring system of naming plumages and molts born to unrelated parents. Inbreeding de- most commonly used by ornithologists I IBA: Important Bird Area. See Important pression occurs because closely related today. In this system, the plumage worn Bird Areas Program. [10·113] parents share more genes, and thus their for the longest time each year (the non- offspring are more likely to receive two breeding plumage), usually produced by ileum: The short, final portion of the small copies (one from each parent) of alleles a complete molt, is called the bird’s basic intestine, running from the jejunum to that cause deleterious traits or genetic plumage, and other plumages are called the large intestine. [4·122] diseases. For example, in humans, the alternate and supplemental. In addition, ilium (plural, ilia): One of the three paired genetic disease hemophilia was once molts are named for the type of plumage bones fused to form the pelvic girdle; the common among some of the inbred royal they produce, not the feathers they shed. ilium forms the cranial and lateral por- families of Eurasia. [9·66] In the traditional system, a bird’s breed- tion, and in birds is completely fused with incest: A mating between close relatives. ing plumage is considered to be the main the ischium and the synsacrum. It has a [6·94] plumage. [3·33, 3·35] cup-shaped depression for the attach- ment of the femur (thigh bone). [4·24] incest taboo: The instinctive avoidance hybridization: Breeding that occurs be- of breeding with close relatives; it has Cornell Laboratory of Ornithology Glossary Inc – Int 37 undoubtedly evolved in many organ- indigobirds: Members, along with whyd- cerned with balance— and the cochlea isms because it reduces inbreeding de- ahs, of the African genus Vidua (mem- and its enclosed cochlear duct, which are pression (see separate entry). Because bers of this genus are called viduines). concerned with hearing. [4·57] determining relatedness can be difficult, Viduines are colorful seed-eaters that inner shell membrane: In avian biology, many species seem to avoid incest by not are brood parasites on other members of the membrane just inside an egg’s outer breeding with any individual with which their family, Estrildidae; they are known shell membrane; it is thinner and less they have been in close contact during for the intricate patterns inside the coarse than the outer membrane. The their developmental period (which is mouths of their nestlings, which strik- inner shell membrane contains the albu- usually a sibling or parent). [10·75] ingly resemble those of the nestlings of men and adheres tightly to the white of incubation: The process by which ani- their hosts. [8·142] a hard-boiled egg, making it difficult to mals that lay external eggs keep those indirect fitness: An individual’s indirect peel. [8·62, 8·69] eggs at the proper temperature for em- fitness is the portion of its genes that is inner vane: The vane located on the bryonic development until they hatch transferred into the next generation (and medial side of a wing or tail feather. On (or the nest fails). Only birds, crocodiles, eventually beyond) as a result of that a wing feather, the inner vane is on the pythons, and monotremes (egg-laying individual’s blood relatives producing edge of the wing that trails in flight. In mammals) incubate their eggs. In most offspring. Some of an individual’s genes flying birds, the inner vane is wider than cases, birds sit on their eggs to keep are propagated when a relative breeds the outer vane, producing an asymmetry them warm, but many megapodes bury because the individual shares a percent- that aids in flight. [3·3] them—in piles of decaying vegetation, age of its genes with that relative (the per- in long tunnels or broad pits where the centage depends on the relatedness), and inner wing: The portion of the wing from earth is warmed from nearby hot streams because the relative passes 50 percent the wrist to the shoulder; the secondary or volcanic cinder fields, or in pits or bur- of its genes (both those that it shares and feathers are located on one section of the rows where bare sand or soil is heated by those that it doesn’t share) to its offspring. inner wing. [5·23] the sun. In very hot environments incu- This contrasts with direct fitness, which innominate arteries: Two large arteries bation may require cooling the eggs by is the portion of genes transferred through (left and right) that branch from the aortic shading them, burying them in sand, or the production of an individual’s own off- arch and then soon branch again into the keeping them moist. [8·93] spring. [6·86] carotid arteries to the head and neck and incubation patch: See brood patch. individual recognition: The ability of an the subclavian arteries to the front limbs [8·94] animal to identify other specific individu- (wings). [4·82, 4·84] incubation period: The time from the als. [7·42] insect-net theory: A variation on the start of regular, uninterrupted incubation inertial navigation: Finding your way by cursorial theory of the origin of avian to hatching. [8·96] keeping track of all the turns and accel- flight, first proposed by John Ostrom in 1976. The insect-net theory suggests that incubation pouch: A type of brood patch, erations you have taken. For example, logging in your brain the turns and ac- dinosaurian bird ancestors first evolved found on the breast of some albatrosses, feathers as thermal insulation; and then, that is a featherless cavity surrounded by celerations of an outward trip and then integrating them to compute a direct as running dinosaurs clapped the feath- thick feathers into which the single egg ered forelimbs together to catch insects, fits so snugly that it may remain inside route home. Inertial navigation requires no outside reference points. [5·82] eventually the motion became flapping even when the bird stands. [8·96] flight. [E·16] indeterminate layers: Bird species that information center hypothesis: The idea that one benefit of colonial breeding is insemination: The transfer of sperm from will lay additional (replacement) eggs the male into the genital tract of the fe- if one or more eggs are removed from that individuals who have been unsuc- cessful at finding food might find better male; in birds, sperm enter the cloaca. the clutch during laying. Once they have [4·133] begun incubation on a full clutch, how- feeding grounds by watching at the col- ever, they cease to lay replacement eggs ony for successful foragers (those who re- insertion: On a skeletal muscle, the end if more are removed. For comparison, see turn with food), and then following them (attachment site) that moves the most determinate layers. [8·90] to good hunting spots. [6·59] during contraction. [4·27] index: A numerical sequence or other infundibulum: The flattened, funnel- inshore feeders: Birds (or other animals) representation that indicates the relative shaped opening of the oviduct. In birds, that forage close to shore, fairly near their level, degree, or amount of something or when the ovary releases an egg, the in- nesting areas; includes birds such as terns some property. For example, an index of fundibulum moves up to the ovary and and many gulls. [6·68] population density, or an index of health. opens, “swallowing” the egg much like a insight learning: A modification of behav- [9·61] snake swallows a rat. [4·130] ior that occurs by evaluation of a situation indicator species: A species that acts innate behavior: See instinctive behav- rather than as a result of previous expe- as a “canary in a coal mine,” because ior. [6·5] rience with a particular problem. [6·14] changes in its population density or dis- innate rhythmicity: The ability to con- instinctive behavior: A behavior that is tribution provide an early warning that its tract without being stimulated by the ner- triggered in full form, without any learn- habitat is changing in some way—often vous system, as found in cardiac muscle ing, the first time an individual responds through degradation by human activity. cells. [4·31] to the releaser; also called innate behav- Birds are good indicators because they inner ear: Complex structure inside the ear ior or instinct. [6·5] are relatively noticeable and easy to sur- in the form of a membranous, fluid-filled integumentary system: The skin and vey, because many bird species are high sac (the membranous labyrinth) floating structures that are produced by the skin, in the food chain and thus sensitive to the inside a bony, fluid-filled sac (the bony such as (in birds) feathers, color pig- bioconcentration (see separate entry) of labyrinth). The inner ear consists of the ments, scales, claws, beak, wattles, and toxins, and because many species have semicircular canals enclosing their ducts comb. [4·3] narrow habitat requirements. Also called and the vestibule enclosing the utriculus intention movements: Movements that biological indicators. [10·108] and the sacculus—all of which are con- are either incomplete (such as the initial Handbook of Bird Biology 38 Int – Kin Glossary stages of a behavior) or that indicate what iridescent colors: Structural colors, some- jugular veins: Paired veins (left and right) the performer is about to do. For example, times brilliant, that shimmer and glitter carrying blood from the head and neck a bird about to attack may crouch down because they change in brightness as the region; they merge on each side with the and tense its muscles. Many intention angle of view changes. The colors are pro- subclavian and pectoral veins to form movements have become exaggerated duced when light waves reflected off thin the cranial vena cavae, which return the and incorporated into displays. [6·30] films (in birds, structural layers within flat- blood to the heart. [4·83] interactive playback experiments: Com- tened feather barbules) interfere with one jugulum: The lower part of the throat of puter-controlled playback experiments in another, as can be seen in soap bubbles birds, just below the gular region. [1·7, which researchers can change various as- and most hummingbirds. [3·54] 1·8] pects of songs played to territorial birds in iris: The colored part of the eye surround- juvenal plumage: Feather coat worn by the middle of a singing encounter, based ing the pupil; it is part of the eye’s choroid juvenile birds after they have molted on the subject bird’s singing behavior. For (middle) layer. The iris contains muscle their natal down; it consists of the first example, researchers might choose to fibers and controls the diameter of the true contour feathers. [3·29] play longer or shorter songs, or songs that pupil and thus the amount of light that match those of the subject, or they might enters the eye. [4·4] juvenile: A young bird. [3·29] respond more slowly or more quickly, or irruptive migration: Migratory move- overlap the subject’s singing. [7·49] ments that are irregular in time and interference: In ecology, a situation in space, depending upon factors other K which one individual actively prevents than a change of seasons, such as food keel: A midventral ridge of bone that proj- other individuals from obtaining a limit- availability. For example, the seeds and ects outward from the sternum and pro- ed resource. It may do so through various buds eaten by finches such as Pine Siskins vides a site for the attachment of the large behaviors, such as aggressive interac- and redpolls fluctuate in abundance not pectoral flight muscles. Large, flightless tions that set up dominance hierarchies only seasonally but from year to year and birds lack a keel, as do the tinamous. Also or territories. [9·42] from region to region, so in some years called a carina. [4·23] International Union for the Conser- large numbers of the birds move out of keratin: A hard protein that forms scales vation of Nature (IUCN): International northern forests to breed, yet in others and claws and is the primary structural nongovernmental organization based they stay put. [5·55] component of mature feathers. Avian ker- in Switzerland and devoted to the con- ischium (plural, ischia): One of the three atin differs from the keratin of all other ani- servation of species. In 1963, the IUCN paired bones fused to form the pelvic mals in its amino acid sequence. [3·28] drafted the original text of the Conven- girdle; the ischium forms the caudal and kettle: A large aggregation of birds, usu- tion on International Trade in Endan- lateral portion. [4·24] ally hawks, that are spiraling upward in gered Species, also known as CITES (see island endemics: Species that evolved on a thermal. [5·39] separate entry), which was finally ratified an island and are found only upon that in 1975. [10·93] keystone species: Species that affect island. [9·55] many other species in their community, interpupillary distance: The distance be- isthmus: The region of the avian oviduct and whose removal would precipitate a tween the centers of an individual’s two after the magnum and before the shell reduction in species diversity or would pupils; important in fitting binoculars. gland; the isthmus secretes the egg and produce other significant changes in [2·30] shell membranes and fluid albumen community structure. [9·126, 9·128] interspecific competition: Direct com- around the fertilized ovum as the ovum kHz: See kilohertz. [7·4] petition for limited resources among in- travels down the oviduct. [4·130] dividuals of different species. [9·56] kidnapping: In avian biology, the ag- IUCN: See International Union for the gressive takeover of a brood of young by intestinal lymph trunk: Lymph vessel that Conservation of Nature. [10·93] adults that are not the parents. [8·128] carries the products of fat digestion from lymph vessels coming from the small kidneys: Paired organs of the excretory intestine to the thoracic duct. These di- system that are irregular in shape and, gestive products are first picked up at J in birds, are each composed of three in- the small intestine by lymph capillaries, jejunum: The long middle portion of the terconnecting lobes; they remove waste which carry them to progressively larger small intestine, running from the duode- products from the blood, especially ni- lymph vessels running along the surface num to the ileum. [4·122] trogenous wastes, and form a highly con- of blood vessels in the intestinal wall. centrated urine that, in birds, is passed to jizz: Birding term for a quick impression the cloaca via the ureters. Kidneys also These vessels eventually join to form the of a bird’s major features. Jizz harkens intestinal lymph trunk, which carries the maintain a balance of salt ions in the back to the “general impression of size blood. [4·125] products to one of the two thoracic ducts, and shape” (G. I. S. S.) that British ob- which eventually enter the cranial vena servers used during World War II to dis- kilohertz (kHz): A measure of the fre- cava. The reason for this circulatory pat- tinguish between enemy and friendly quency of a sound, in thousands of cycles tern, which occurs in (at least) all birds aircraft. [1·24] (of compression and thinning of the air) and mammals, is not understood. [4·89] per second. One kilohertz equals 1,000 journal: A written record of field observa- Hertz. Also written kiloHertz. [7·4] intraspecific competition: Competition tions of birds, other animals, or natural (for food, territories, mates, and so on) phenomena. [2·47] kingdom: Level of classification of or- that occurs among members of the same ganisms above “phylum”; similar phyla species. [9·39] jugal arch: Bony rod on each side of the are placed within the same kingdom. upper jaw below the palatine (another ioras: Small, arboreal songbirds that All birds are in the kingdom Animalia. set of bony rods). In birds, as the lower [1·52] search leaves, often in dense foliage, for jaw opens it moves the quadrate, which insects. The four species form the family pushes the palatine and jugal arch for- kin selection theory: A line of reasoning Aegithinidae, endemic to the Oriental ward, pushing on the premaxillary bones suggesting that the closer the kinship (de- zoogeographic region. [1·89] to raise the upper jaw. [4·12] gree of relatedness) between two animals Cornell Laboratory of Ornithology Glossary Kiw – Lim 39 of the same species, the greater will be it acts as a valve, regulating the flow of air copulations with visiting females. In most their tendency to cooperate with one an- into the trachea. In mammals the larynx lekking species, the top few males secure other in various ways. Such cooperation is the voice box, but in birds it produces most of the matings, resulting in a high results from natural selection because no sound, leaving sound production to degree of polygyny. [6·75] if an animal enhances the survival of a the syrinx. [4·91] lens: Spherical or ovoid structure near relative, it also enhances its own indirect lateral: Toward the side of the body; away the front of the eye; it changes its curva- fitness (see separate entry); the closer the from the midline. [1·4] ture to sharply focus images from varying relative, the greater the degree to which lateral labia: Important sound-producing distances on the retina. The lens is crys- its own indirect fitness is enhanced. talline-like and composed of regularly [6·93] membranes in the lateral wall of each half of the syrinx. During sound production oriented layers of collagen fibers. [4·47] kiwi: A family (Apterygidae) of three spe- muscles move the bronchi upward into the Lepidosauromorpha: One of the two ma- cies of grouse-sized, flightless, nocturnal trachea, twisting the third bronchial car- jor groups of diapsid reptiles; it contains ratites endemic to New Zealand; kiwis tilages so that the lateral labia and medial all snakes and lizards as well as the an- probe soil with their long beaks, using labia (in the medial walls) move toward cient ichthyosaurs and plesiosaurs. [E·8] their keen sense of smell to locate earth- (and close to) each other and into the path worms. [1·96] lesser coverts: The feathers on the upper of air flowing out of the respiratory system. surface of the wing that partly overlie the koilin: See gastric cuticle. [4·119] Sound is produced as air rushes between median coverts and extend to the mar- K-selected species: Species in which the lateral and medial labia, causing these ginal coverts. [1·12, 1·13] soft tissues to vibrate. [4·96] individuals have a life history strategy leukocytes: See white blood cells. Also that relies on longevity, rather than a Laurasia: The northern land mass formed spelled leucocytes. [4·88] high reproductive rate, to maximize the 200 million years ago when Pangea split number of offspring produced during into two large land masses. It consisted liana: A woody vine. [9·88] their lifetimes. Such species tend to be of present-day North America, Europe, life history theory: A set of ideas that large, develop slowly, begin breeding at and Asia. [1·68] attempt to explain the diversity in the a relatively old age, have few young per leafbirds: Eight species of oriole-sized breeding strategies of living things by cycle (small clutches or litters), breed arboreal songbirds endemic to the Ori- looking at why various characteristics infrequently, take care of their young ental zoogeographic region; leafbirds relating to an organism’s birth, death, for extended periods, and have long life are mostly green and yellow, and feed and reproduction have evolved. These spans. [9·45] mainly on insects and fruit. Together with characteristics, called life history traits K·T event: The massive, worldwide ex- fairy-bluebirds, they form the family Ire- (see separate entry), include such things tinctions 65 million years ago at the end nidae. [1·88] as number of offspring, age at first breed- of the Cretaceous Period and the begin- ing, and length of the developmental leaf tossing: A foraging technique in period. [8·3] ning of the Tertiary Period, which wiped which a bird tosses aside the leaf litter out dinosaurs as well as many other (with its beak or feet) to search for food; life history traits: Characteristics of liv- plants and animals. The extinctions are used by towhees, turkeys, thrashers, and ing things that are related to birth, death, thought to have resulted from worldwide many sparrows. [6·47] and reproduction. Examples include the climatic disturbance caused by a large number of offspring, the age at first breed- meteor colliding with Earth. [E·25] learned behavior: A behavior that re- ing, the interval between breeding cycles, quires some amount of previous experi- and the chance of surviving to various ence—such as exposure to a stimulus— ages. Collectively, these characteristics as well as a memory of that experience, make up an organism’s life history or life L before it is carried out fully. [6·9] history strategy. [8·3] lacrimal gland: One of the two tear learning: The modification of a behavior life list: Record of every species seen by a glands on each side of the eye; the lac- as a result of experience. [6·8] particular person, noting the date and loca- rimal gland lies in the lower part of the leg spurs: Bony outgrowths near the dis- tion of the first sighting. People may keep life orbit of the eye and has many ducts enter- tal end of the tarsometatarsus, covered by lists of different groups of organisms, such as ing the space between the lower lid and a pointed horny sheath; used as weapons birds, butterflies, or reptiles. [2·54] the cornea. Its secretions moisten the eye by male chickens, peafowl, and many and nourish the cornea. [4·46] lift: Force acting on a moving airfoil (such other pheasant relatives. [4·26] as a bird’s wing) perpendicular to the di- laminar flow: A smooth flow of air over lek: 1. A traditional courtship area where rection of airflow; in a bird that is gliding an airfoil. [5·13] many males of the same species gather or flying horizontally, lift acts upward in large intestine: Short, straight tube extend- to attract females for mating; each male opposition to gravity. [5·10, 5·13] ing from the small intestine to (in birds) the spends a large amount of time defending lift-to-drag ratio: The lift produced by a cloaca; it holds the intestinal contents while a small site at which he displays to com- wing divided by the drag it experiences water is being reabsorbed, and passes the pete with other males and, in particular, while flying. Long, narrow wings have remainder to the cloaca. [4·123] to earn copulations with sexually recep- the greatest lift-to-drag ratios and are the laryngeal cartilages: Two major cartilages tive females who visit the lek to choose most efficient. [5·35] that make up the sides and floor of the among the males. In most species, the top few males secure most of the matings. ligament: Fibrous connective tissue that larynx. Also called the cricoid cartilages. connects one bone to another across a [4·91] The lek contains no nest sites, food, or other resources useful to nesting females. joint. [4·6] laryngeal folds: Fleshy folds in the lower [6·75] 2. The group of males gathered at limiting factor: Something that is present surface of the pharynx that surround the traditional courtship site described in in (or missing from) a particular environ- the glottis, the opening to the larynx. definition 1. [6·75] ment and as a result prevents a particular [4·112] lek polygyny: Mating system in which species from living or breeding in that larynx: Structure at the upper end of the males gather in leks and display to earn place. Examples of limiting factors in- trachea that consists of several cartilages; clude a large number of predators and Handbook of Bird Biology 40 Liv – Mat Glossary a low level of a critical resource, such as oxygen and nutrients to the body (tis- the fertilized ovum as it travels down the food. [9·10] sue) cells and picked up carbon dioxide oviduct. [4·130] liver: The largest internal organ in the and other wastes diffuses into the lym- malar region: Small area caudal to the body; in birds the liver has two lobes and phatic capillaries, and is then termed base of the lower beak; also called the performs a variety of functions, includ- lymph. The lymphatic capillaries lead to cheek. [1·7, 1·8] progressively larger lymphatic vessels, ing secreting bile to help emulsify fats for malar stripe: A distinctively colored digestion, storing sugars and fats, form- which eventually dump their contents into the venous system. [4·88] stripe in the malar region of birds; also ing uric acid, and removing foreign sub- called a mustache stripe or whisker stances from the blood. [4·124] lymphatic system: Organ system—com- stripe. [1·8, 2·10] posed of lymphatic vessels, ducts, and lithornithids: Extinct, flying, chicken-like manakins: A Neotropical suboscine fam- palaeognathous birds thought to have nodes—that gathers tissue fluid that has leaked from the blood capillaries, filters ily (Pipridae, 44 species) of small, fru- been common in North America and Eu- givorous birds noted for the stereotyped rope during the early Tertiary; they may foreign substances and old or damaged cells from the fluid, and then returns the courtship displays of the colorful males. be the ancestral stock that gave rise to Many manakin species form leks, and ratites all over the world. [E·23] fluid to the general blood circulation. The lymphatic system also releases antibod- most displays involve intricate maneu- local population: A population confined ies and transports the products of fat di- vers that may be enhanced by mechani- to a small area; for example, all the Blue gestion from the intestines to the venous cal and/or vocal sounds and plumage Jays in one neighborhood. [9·49] system, bypassing the liver. [4·88] displays. [6·78] lore: Small area between the eye and the lyrebirds: Two pheasant-sized songbirds mandible: In birds, usually refers to the base of the upper beak. [1·7, 1·8] (forming the family Menuridae) of Austra- lower half of the beak; called the lower lower beak: The lower half of the beak; lian rain forests, named for the elaborate, beak or lower jaw in the Handbook of Bird sometimes called the mandible or lower harp-shaped tail of the Superb Lyrebird. Biology. [4·11] jaw. [1·6] They have loud songs and calls and are mandibular nerve: A division of the tri- lower critical temperature: The envi- fantastic mimics, sometimes clearly re- geminal nerve (the fifth cranial nerve) ronmental temperature below which producing mechanical sounds, such as after it leaves the brain; the mandibular the body of a bird or mammal increases logging trucks and chain saws, along nerve carries sensory input from the low- its metabolic rate and employs physio- with the songs of other birds. [1·94] er beak and corner of the mouth, and mo- logical responses to warm itself, assum- tor output from the brain to the muscles ing behavioral responses are no longer of the lower beak. [4·41] adequate. [9·16] manus: The portion of the forelimb distal lower jaw: See lower beak. [1·6] M to the wrist; also called the hand. The pri- macroevolution: The evolution of new mary feathers attach to the manus. [1·9] lower lethal temperature: The environ- species over long periods of time, such mental temperature below which a bird as thousands of years (see microevolu- map and compass model: The theory or mammal cannot keep its body warm tion). [1·37] (proposed by Gustav Kramer in 1953) enough to survive. [9·16] that two things are required to navigate magnetic anomaly: A place where the to a particular destination from an unfa- low pressure system: A weather system earth’s magnetic field is disturbed, usu- miliar location: (1) a map indicating cur- consisting of cold and warm air masses ally by large deposits of iron near the rent location with respect to destination, circulating around an area of low baro- surface. [5·98] and (2) a compass indicating the desired metric pressure. In the Northern Hemi- magnetic compass: A mechanism by direction of travel. [5·84] sphere the masses circulate counter- which birds and some other animals can marginal coverts: The feathers covering clockwise; in the Southern Hemisphere use the earth’s magnetic field to deter- they move clockwise. [5·69] the upper surface of the wing, from the mine compass direction. [5·89] leading edge back to the greater coverts, lumbar vertebrae: The vertebrae of the magnetic field of the earth: The region which they partly overlie. [1·12, 1·13] lower back; in birds they are all fused around the earth in which objects experi- with the sacral vertebrae and some of the market hunting: Intensive hunting of spe- ence magnetic force (which is a vector—a cies to obtain meat, feathers, hides, or thoracic and caudal vertebrae to form the quantity with both strength and direction). synsacrum. [4·15, 4·18] other body parts highly prized by com- The field is created by currents generated mercial consumers. [10·11] lumbosacral enlargement: Swelling by the constant motion of molten iron in along the spinal cord at the level of the the earth’s core. In effect, the earth is a marine birds: See seabirds. [1·65] legs; it is associated with the lumbosacral large magnet with two magnetic poles matched countersinging: Interaction in plexus. [4·40] (near the North and South Pole). The field which two birds sing back and forth to lumbosacral plexus: Plexus (see separate is stronger closer to the magnetic poles one another, each choosing songs that entry) along the spinal cord at the level of and weaker toward the equator. [5·90] are identical or similar to those of the the hind limbs; it is associated with the magnetic map: A mechanism by which other, or that contain similar phrases. lumbosacral enlargement of the spinal birds (and some other animals) may be Not well understood, but thought to be cord. [4·39] able to use the earth’s magnetic field as a one way in which birds “duel” through song. [7·36] lymph: Fluid carried in lymph vessels; map. For example, because the magnetic also see lymphatic capillaries and lym- field is strongest toward the poles, a bird matrilineal societies: Groups of animals phatic system. [4·87] might use the strength of the field at any in which the females remain in their na- given point to estimate latitude. [5·97] tal colonies but the males disperse when lymphatic capillaries: Tiny vessels that they become adults. Thus, within a colo- form a network intertwining with the magnum: The first region of the avian oviduct, after the infundibulum; the mag- ny, the females share more genes than do capillary beds of the arterial and venous the males, and are more likely than males systems throughout most body tissues. num is glandular and secretes the first of the albumen or “white” of the egg around to engage in acts that appear altruistic Some of the tissue fluid that has lost its toward one another. [6·56] Cornell Laboratory of Ornithology Glossary Max – Moa 41 maxillary nerve: A division of the tri- mesencephalon: See midbrain. [4·38] [4·38] geminal nerve (the fifth cranial nerve) as mesites: A family (Mesitornithidae, 3 middle ear: Air-filled chamber between it leaves the brain. It carries sensory input species) of rail-like, ground-dwelling the eardrum and the cochlea (inner ear); from the skin of the face, upper jaw, up- birds endemic to Madagascar; mesites in birds it houses the columella. [4·57] per eyelid, and conjunctiva (tissue cover- are about the size of a Mourning Dove. ing the eye). [4·41] mid-story vegetation: Layer of vegetation [1·87] in a forest that consists of trees whose medial: Toward the midline of the body. mesobronchus: The continuation of crowns are below the level of the main [1·4] each bronchus (right and left) as it enters canopy and above the understory vege- medial labia: Important sound-pro- the lung and loses its cartilaginous half- tation (see separate entry). [9·93] ducing membranes in the medial wall rings; also called the primary bronchus. migration: The regular movement of all of each half of the syrinx. During sound The mesobronchi gradually decrease or part of a population to and from an production muscles move the bronchi in diameter and branch into secondary area; usually refers to seasonal journeys upward into the trachea, pushing the me- bronchi. Only in birds is the term “meso- to and from breeding grounds or feeding dial labia and lateral labia (in the lateral bronchus” used as an alternate name for areas. [5·52] walls) toward (and close to) each other the primary bronchus. [4·98] and into the path of air flowing out of the migratory program: The genetically metabolic water: Water obtained by an programmed information that guides an respiratory system. Sound is produced as organism’s body as a byproduct of the air rushes between the lateral and medial inexperienced bird on its first migration. chemical breakdown of fats, carbohy- The timing of that migration is controlled labia, causing these soft tissues to vibrate. drates, and proteins within the body. [4·96] by an internal biological clock that con- [9·20] trols circannual rhythms, and the bird median: On the midline of the body. metabolism: All the chemical processes chooses the approximate direction and [1·4] that take place in the cells and tissues of distance through its built-in ability to median coverts: The covert feathers lying the body. [4·144] carry out vector navigation. [5·79] between the lesser and greater coverts metacarpals: Palm bones. In humans migratory restlessness: Nocturnal hop- on the upper surface of the wing. [1·12, they remain distinct, but in birds, they ping and fluttering during the normal 1·13] are fused with some of the carpals (wrist migratory period, performed by caged medulla oblongata: The most posterior bones) to form the large carpometacar- birds that normally are inactive at night. portion of the brain, also called the brain pus. [4·21] Researchers often use the degree of this stem, where the nuclei of most of the metapopulation: All the individuals of a “unrest” as an indication of a bird’s de- cranial nerves are located. The medulla species living in an area that contains a sire to migrate, and the orientation of oblongata extends caudally through the set of subpopulations (local populations) the hopping and fluttering to indicate foramen magnum to become the spinal close enough together so that individuals the compass direction in which the bird cord. [4·37] from occupied habitat patches can dis- would normally be migrating. First called megapodes: A family (Megapodiidae, 21 perse and recolonize patches where the Zugunruhe by the German scientists who species) of chicken-like terrestrial birds species has gone extinct. [10·79] discovered it. [5·60] of the Australasian and Oriental regions. metapopulation dynamics: The changes mimicry: A situation in which one indi- Megapodes do not use their own body in local populations found within a meta- vidual or species has evolved or learned heat to incubate their eggs; instead, in population over time, in which the local to be similar to another in appearance, many species the males tend the eggs of populations (in a habitat patch) may go behavior, or sound. For example, Brown several females in huge, warm mounds extinct and then individuals from within Thrashers and Northern Mockingbirds of decaying vegetation. Other species the metapopulation may recolonize the imitate the songs of other bird species. use geothermal heat to warm their eggs. patch at a later date, producing a long- [7·81] [1·93, 6·70] term situation in which local populations mineral lick: General term for a spot that melanin: Pigment, usually present as tiny “wink” out and then “wink” back on. animals visit to obtain minerals, either granules, that produces a range of earthy [10·77] naturally occurring in the soil or provided colors from dark black, brown, and red- metapopulation models: See population by humans. A clay lick (see separate en- brown to gray, yellow-brown and pale viability analysis. [10·77] try) is one type of mineral lick. [9·28] yellow. Birds can synthesize their own metatarsals: Instep bones. In humans mitigation bank: See wetland mitigation. melanin by oxidizing the amino acid ty- [10·93] rosine. [3·50] they remain distinct, but in birds they fuse with the distal tarsals (ankle bones) mitochondria: Membrane-bounded membranous labyrinth: A system of in- to form the tarsometatarsus, the long units in all cells except bacteria; they terconnected, fluid-filled canals (con- bone supporting the upper section of the generate most of the cell’s energy. Mito- taining endolymph) floating in the bony foot. [1·14, 4·25] chondria contain DNA in the form of a labyrinth of the inner ear. The labyrinth ring. [1·44] forms the cochlear duct, the semicircu- metatarsus: See tarsometatarsus. [1·14] lar ducts, and the utriculus and sacculus. microevolution: A change in the frequen- mnemonic device: A memory aid. In [4·57] cy of a genetically controlled characteris- birding, usually refers to the association of phrases from human speech with the meninges: General term for the vascu- tic in a population over a relatively short period of time (see macroevolution). songs of particular birds, to help people larized membranes surrounding the remember the songs more easily. [2·16] brain and spinal cord. Specifically, these [1·37] are the outer fibrous dura and the inner midbrain: The middle region of the ver- moas: A family (Dinornithidae) of at least arachnoid and pia layers. Meninges pro- tebrate brain; it contains the optic lobes 22 species of enormous, flightless ratites vide sustenance and waste removal for (which in birds are huge and dominant), that roamed the open foothills and tus- the cells of the brain and spinal cord, as well as regions for the input and pro- sock lands of interior New Zealand. Moas which are not served by the circulatory cessing of information on hearing and evolved in the late Tertiary, but were extir- system. [4·36] balance. Also called the mesencephalon. pated by the Polynesians approximately Handbook of Bird Biology 42 Mob – Neo Glossary 400 years ago. The largest moa stood 14 shaped, mud nests. The Magpie-lark is found in muscle cells. Similar in both feet tall. [1·96, E·24] widespread, abundant, and well-known form and function to the hemoglobin mobbing: Behavior in which a number of throughout open areas in much of Austra- in blood cells, myoglobin binds with birds (often different species) swoop and lia; the Torrent-lark inhabits fast-flowing oxygen, storing it until the muscle cell dash at a potential predator; they usually streams in the mountainous areas of New needs it to release energy. [5·7] give broad-band, raspy calls (mobbing Guinea. [1·96] myopia: Visual condition of an indi- calls) that are easy to locate and thus at- muscle fibers: Bundles of long, cylin- vidual in which distant objects blur be- tract additional birds. [2·26, 6·51] drical, contractile cells (those that can cause images are focussed in front of, model song: A song that a bird listens to contract) that are the functional units of rather than on, the retina; also called and attempts to duplicate during the pro- muscle tissue; they shorten when stimu- nearsightedness. Some types of birds, cess of song learning. [7·26] lated by a nerve impulse. [4·26] such as penguins, are myopic on land mustache stripe: A distinctively colored because their eye is designed for vision molting: The process of shedding all or under water. [4·50] part of the feather coat and replacing it stripe in the malar region of birds; also with new growth. [3·28] Note: For spe- called a malar stripe or whisker stripe. cific names of molts, see p. 3·35. [1·8, 2·10] monocular vision: Type of vision that mutation: A change in the sequence of ni- N produces flat, two-dimensional images, trogenous bases in DNA. Most mutations nape: The back of the neck. [1·7] have little or no effect on an organism, but in contrast to binocular vision, which naris (plural, nares): The openings of the a few result in changes in the structure or produces three-dimensional images. nasal cavity; they are located in the upper type of proteins produced, thus creating Monocular vision results when the eyes beak, usually near its base. Also called the genetic diversity upon which natural are positioned on the sides of the head nostrils. [3·40] such that an object can be seen by one selection can act. [1·36, 1·41] nasal-frontal hinge: See craniofacial eye or the other, but not by both eyes at mutual displays: Intricate, synchronized hinge. [4·11] the same time. [4·51] displays or dances performed by mem- nasal region: Portion of the skull contain- monogamy: Mating system in which one bers of a mated pair; the displays appear ing the nostrils. [4·11] male pairs with one female, at least for a to stimulate and coordinate breeding given breeding season. [6·68] behavior between pair members and to natal dispersal: The movement of a young reaffirm the pair bond. Given by many animal away from the area in which it was morph: A set of individuals within a spe- long-lived birds that mate for life, such born and raised. In birds, natal dispersal cies that are similar to one another in as albatrosses, gannets, grebes, and pen- usually occurs between fledging and the some genetically determined morpho- guins. [6·38] first breeding season. [9·65] logical characteristic, but are distinctly different from other sets of individuals mutualism: An association between two natal down: The soft down feathers cov- within that species. Morphs may differ in (or more) organisms in which both (or all) ering young birds before they molt into characteristics such as color, body size, organisms benefit. [9·7] juvenal plumage. [3·16] or bill length or shape, but not in charac- mycoplasmal conjunctivitis: A highly natural selection: The process by which teristics that are related to sex, age, local- contagious infection of the conjunc- evolution occurs: as individuals with ity, or season. [9·42] tiva of the eyes that has spread rapidly traits that allow them to compete better mortality rate: See death rate. [9·49] through House Finch populations in east- for essential resources survive and repro- ern North America since its discovery in duce better than other members of their motor neurons: Nerve cells that con- Maryland in 1994. The disease also af- population, they contribute more of their vey impulses from the brain and spinal fects Purple Finches, American Gold- genes to the next generation, and these cord to stimulate a muscle to contract or finches, Evening Grosbeaks, and Pine favorable (adaptive) traits increase in the permit it to relax, or to cause a gland to Grosbeaks, although to a far lesser ex- population. [1·34] secrete. [4·33] tent. Many birds that contract the disease Nearctic region: Zoogeographic region mound nest: A nest composed of a pile eventually die—usually from starvation, including arctic, temperate, and sub- of material with an egg chamber in the predation, exposure to the elements, or tropical North America, reaching south middle; in some species the chamber is some other factor that results from hav- to the northern border of tropical rain entered through a tunnel from the out- ing impaired eyesight. The disease is forest in Mexico. In the Handbook of Bird side. Mound nests are built by many caused by a previously unknown strain Biology Greenland is included in the Ne- megapodes, which bury their eggs in a of the bacterium Mycoplasma gallisepti- arctic, but some sources place it in the pile of decomposing organic material; cum; other strains had long been known Palearctic. [1·70, 1·71] the Hamerkop of Africa, which builds a in poultry. Also called House Finch eye massive mound of sticks; and Monk Par- disease. [9·71] neighbor recognition: In avian biology, the ability of territorial birds to identify akeets and Palmchats, which build large, myelin: Fatty material forming the nerve colonial mound nests. [8·39] their neighbors—usually on the basis of sheath around many axons. The process song alone. [7·43] mousebirds: Long-tailed African birds by which myelin is deposited in a devel- (family Coliidae, 6 species), about the oping embryo and young animal is called Neognathae: One of the two Superor- size of a Mourning Dove, that climb myelination. [4·33] ders of ornithurine birds; it contains all birds with a neognathous palate—one through vegetation using their stout, myelination: See myelin. [4·33] hooked beaks; frequently mousebirds in which the vomer and basipterygoid perch in a “chin-up position,” hanging myelin sheath: The pale, fatty wrapping process are reduced and a flexible joint vertically from a branch. Also known as that surrounds the axons of many neu- exists between the pterygoid and pala- colies. [1·85] rons; it insulates the axon, functioning, tine bones (as compared to a palaeogna- in part, like the insulation around house- thous palate, which is formed by larger, mudnest builders: A family (Grallinidae) hold wiring. [4·33] more rigid bones). Neognathae includes of two striking, black-and-white, robin- all modern birds except ratites, as well as sized birds named for their large, cup- myoglobin: A red-pigmented protein

Cornell Laboratory of Ornithology Glossary Neo – Off 43 the extinct Diatryma and dromornithids. nearly tailless, primarily insectivorous northeaster: Winter storm that sweeps [E·23] suboscine birds endemic to New Zea- into New England across the Atlantic neognathous: See Neognathae. [E·23] land. One of the four known species, the Ocean from the northeast, the high winds Stephens Island Wren, became extinct in circulating around a low pressure area. neomorphine cuckoo: See cuckoo. the late 1800s. [1·96] Often northeasters push pelagic birds [8·141] NGO: See nongovernmental organiz- from the open ocean near or over land; neoteny: An evolutionary phenomenon ation. [10·115] in severe storms, many birds die. Also in which juvenile traits are retained into called nor’easter. [5·72] adulthood. [5·51] niche: The role played by a particular species in its environment. Niche in- Northern Marine Region: The major fau- Neotropical migrants: Birds that winter cludes the many ways in which a species nal region of the seas that includes the in the Neotropics but migrate to the Ne- interacts with its physical and biotic en- frigid waters of the Arctic south to about arctic region to breed. Examples include vironment, such as what and how it eats, 35 degrees north latitude. [1·99] many wood-warblers, tanagers, and ori- what temperatures it requires, where it northern timberline: See arctic tundra/ oles. [1·71] spends time, when it is active, whether coniferous forest ecotone. [9·123] Neotropical region: Zoogeographic re- it disperses seeds or pollinates plants (if notched tail: See emarginate. [1·19] gion including the West Indies, South an animal), what animals pollinate it (if a America, and Central America north to plant), what preys on it, and so on. Also nucleus (plural, nuclei): 1. The mem- the northern edge of the tropical forests called ecological niche. [9·102] brane-bounded command center of all cells except bacteria; usually the nucleus in Mexico. Also called the Neotropics. niche shifting: Describes a situation in [1·70, 1·72] contains the chromosomes. [1·44] 2. which a species occupies a different Clusters of nerve cell bodies within the nerve: A bundle of many nerve cell fibers, niche in different communities, depend- central nervous system. Collectively nu- surrounded and bound together by con- ing on which competitors are found in clei form the gray matter, found at the nective tissue; one nerve is large enough each. [9·103] core of the spinal cord and in the outer to be seen with the naked eye. [4·32] nictitating membrane: A thin, translucent areas of the brain. [4·36] nerve cell: See neuron. [4·32] fold of skin that sweeps sideways across nuptial plumage: See alternate plumage. nest: In avian biology, a structure built, the eye from front to back, moistening [3·33] excavated, or taken over by a bird, in and cleaning the eye and protecting its which the eggs are laid and remain until surface. Found in many vertebrates, in- they hatch. In many species, the young cluding birds, but not in humans. [1·7] remain in the nest until they are able to night flight calls: Calls given by migrating O fly. In some species, the “nest” is simply a birds as they fly at night. [7·44] objective lens: The large lens in a pair of binoculars or a telescope that is far- scrape or depression on the ground. See nocturnal: Active at night. [9·31] specific nest types, such as cup nest, for thest from the eye; it receives the image more information. [8·18] node: A clump or mass of tissue, for ex- viewed in the eyepiece. [2·35] ample lymph nodes [4·88] or the nodes obligate brood parasites: Bird species that nest appropriation: Nesting in a nest that in the heart muscle that stimulate the always lay their eggs in the nests of other previously was used by another species heartbeat. [4·79] or another member of the same spe- species, leaving the resulting young to be cies—usually after the previous breeding nomadic: Pattern of movement in which raised entirely by the host parents. For attempt has ended. Nest appropriators individuals are constantly on the move, comparison, see brood parasite. [8·139] showing no tendency to return to previ- include cavity adopters as well as spe- obligate partial migrant: See partial mi- ously occupied places. Crossbills and cies that take over open nests, such as gration. [5·56] Solitary Sandpipers, Bonaparte’s Gulls, perhaps Budgerigars may be considered obligatory annual migration: A type of House Sparrows, Great Horned Owls, nomadic. [5·56] migration in which all individuals of a Little Swifts, and many others. [8·58] nonbreeding plumage: See basic plum- species migrate to and from a particular age. [3·33] net primary productivity: See primary area each year. Usually obligatory migra- productivity. [9·87] nongovernmental organization (NGO): tion occurs in species whose breeding- neuroglia: Cells that form a supporting, A nonprofit organization that is indepen- area resources vary greatly from season protective, felt-like bed for neurons and dent from the government. [10·115] to season in a predictable way—for ex- also provide electrical insulation. Not noniridescent structural colors: Colors ample, in birds that eat insects or nectar nerve cells themselves, they also are produced in birds when tiny vacuoles and that nest at high latitudes where both called glia or glial cells. [4·36] (pockets) of air within cells in the feather insects and nectar become scarce or ab- neuron: A nerve cell. The basic unit of barbs scatter incoming light. All blues and sent during winter. [5·54] the nervous system, it consists of a cell whites of birds are noniridescent structural occipital condyle: A prominent “bump” body, axon, and one or more dendrites. It colors, as are many greens. [3·55] or peg on the base of the skull, with which is capable of generating, conducting, and North American Waterfowl Manage- the atlas (the first cervical vertebra) ar- receiving nerve impulses. [4·32] ment Plan: A 1986 agreement between ticulates. Birds have one occipital con- New World cuckoo: See cuckoo. the United States and Canada to coop- dyle, but mammals have two. [4·16] [8·141] eratively protect waterfowl by jointly oculomotor nerve: The third cranial protecting habitat, restoring declining nerve. It controls eye movement by car- New World warblers: A family (Parulidae) species, and conducting population of 115 species of small, insectivorous birds, rying motor output to the eye muscles; it research. Each year biologists make de- also carries motor signals to the eyelid many of which are colorful, found in the tailed population estimates, and then New World. Many are Neotropical mi- muscles and the tear gland of the nictitat- use them to help regulate the number ing membrane. [4·41] grants. Also called wood-warblers. [1·71] of individuals of each waterfowl species offshore feeders: Birds (or other animals) New Zealand Wrens: A family (Acanthi- harvested during the waterfowl hunting that hunt schooling fish far out to sea and sittidae) of tiny (smaller than a kinglet), season. [10·97] Handbook of Bird Biology 44 Oil – Ost Glossary far from their nests; includes most pelagic Found in some ground-feeding birds organ: A group of tissues, often of dis- species such as albatrosses and gannets. such as starlings, pigeons, and domestic tinctive character and function, that ag- Avian offshore feeders are also called pe- chickens. [3·40] gregate to form a discrete structure with lagic feeders. [6·68] ophthalmic nerve: A division of the tri- a particular function, such as the heart, Oilbird: A large, nocturnal frugivore of geminal nerve (the fifth cranial nerve) as stomach, or lung. [4·2] South America, related to nighthawks; it leaves the brain; the ophthalmic carries organ system: A group of organs whose the Oilbird is the only member of its fam- sensory input from the nasal cavity (for various functions are coordinated to ac- ily, Steatornithidae. Oilbirds use echo- nonolfactory nasal sensations), eyeball complish one or more of the basic func- location to reach their nests, which are (for nonvisual eye sensations), upper eye- tions of life; examples include the diges- located deep within caves. [4·63] lid, forehead, and upper beak. In ducks tive system and the respiratory system. oil gland: A gland, located at the base and geese, it carries sensory input from [4·2] of the tail on the dorsal side of the bird’s the bill tip organ. [4·41] Oriental region: Zoogeographic region body, that secretes oils that birds spread opposite birds: A group of small to me- including all of Asia south and east of the over their feathers during preening. The dium birds that lived in the Cretaceous Himalayan Mountains (India and South- oils keep the skin supple and the feath- period between 65 and 140 million years east Asia), as well as southern China and ers and scales from becoming brittle, ago. They are called “opposite” birds be- the islands of Indonesia and the Philip- but they do not appear to waterproof the cause their metatarsals (the instep bones pines, east to include the islands of Timor feathers. Also called the uropygial gland of humans) fuse to form part of the tarso- and Sulawesi. [1·70, 1·88] or preen gland. [3·20] metatarsus from the proximal end to the origin: The end (attachment site) of a skel- old-growth forests: Virgin (uncut) forests distal end, a direction opposite to that etal muscle that moves the least during or forests that have remained uncut for of modern birds. Also called enantiorni- contraction. [4·27] thines. [E·20] a very long time and thus contain trees ornithischian dinosaurs: One of the two ranging from hundreds to thousands of optic chiasma: Site at which the optic tract major groups of dinosaurs, called ornith- years in age. [10·22] coming from the left eye crosses the tract ischian (bird-hipped) dinosaurs because Old World cuckoo: See cuckoo. [8·141] coming from the right eye just before en- their hips superficially resembled those tering the optic lobes of the brain. [4·41] olfaction: The sense of smell. [4·62] of modern birds. They were highly spe- optic lobes: Two large lobes of the mid- cialized herbivores. [E·8, E·32] olfactory: Relating to the sense of smell. brain; in birds, they dominate the mid- [4·62] ornithopters: Early flying machines brain and are proportionally larger than whose crude wings were lifted by hu- olfactory epithelium: The lining or sur- those of mammals. The optic lobes re- mans flapping their arms; they never got face tissue of the nasal cavities; it con- ceive the optic tracts from the eyes and off the ground. [5·8] tains the sensory endings of the olfactory are the sites of much initial processing of nerves, which carry input about odors to visual information. [4·37] Ornithurae: One of the two major sub- the brain. [4·62] classes of birds; it includes all modern optic nerve: See optic tract. [4·41] birds as well as the Lithornithids, Am- olfactory lobes: Two lobes at the anterior optic tract: Bundle of sensory nerves car- biortiformes, Hesperornithiformes, and ends of the cerebral hemispheres of the rying visual sensations from the retina of Ichthyornithiformes. [E·20] brain that are concerned with the sense of the eye to the optic lobe on the oppo- smell. They are relatively small in birds. ornithurine: Describes birds in the Sub- site side of the brain. This nerve cable class Ornithurae. [E·20] [4·37] is considered the second cranial nerve, olfactory map hypothesis: The idea that although it is really a tract. [4·41] oscillogram: A visual representation of a sound, plotted on a graph as relative homing pigeons may learn a gradient odor optimal foraging: A feeding strategy that map of the vicinity of their home loft by loudness (vertical axis) versus time (hori- maximizes caloric intake while minimiz- zontal axis). The vertical axis actually is associating airborne odors with the direc- ing the costs of obtaining food, such as tions from which winds carry them past a measure of the increase or decrease in those associated with searching for, cap- air pressure (measured in micropascals) the loft. This gradient odor map would be turing, and handling prey. [9·35] based on small but systematic changes in associated with the sound wave, which the intensity or composition of odors over oral flanges: Brightly colored enlarge- determines the loudness. The oscillogram a large area; as a bird moves in a given di- ments around the base of the bill in nest- does not show sound frequency. [7·7] rection, particular odors become steadily lings of many species in which the parents oscines: Members of Suborder Passeri, stronger or weaker. [5·93] feed the young. The flanges extend from which is one of the two large suborders the corner of the mouth and taper toward of Order Passeriformes (perching birds); olfactory nerve: The first cranial nerve; the tip of the bill, and are well-supplied it carries the sensations of smell from the oscines also are known as songbirds or with tactile nerve endings. Touching a true songbirds. Oscines have particularly lining of the nasal cavity to the olfactory flange causes the mouth to spring open, bulb of the brain. [4·40] complex voice boxes, which allow them and the colors may help parents to place to sing more complex songs than other omnivores: Organisms that eat both the food properly. [3·43, 8·107] birds. [7·25] plants and animals. [9·123] orbit: Cavity in the skull that houses the ossification: Hardening or calcification oölogy: The study of birds’ eggs. [8·80] eye. The eyes of most birds are so large of soft tissue (such as cartilage or tendon) operational sex ratio: The ratio of fertiliz- that the left and right orbits nearly meet at into bone or a bone-like material. [4·6] the midline of the skull. [4·11, 4·13] able females to sexually active males in Ostrich: The largest living bird and the a given population. Thus, the ratio mea- order: Level of classification of organ- only member of its order and family, Stru- sures not just the breeders, but all indi- isms above “family” and below “class”; thionidae. This familiar, flightless ratite of viduals physiologically and behaviorally similar families are placed in the same African deserts and savannas is almost capable of breeding. [9·80] order. The scientific names of bird orders entirely herbivorous. Ostriches breed operculum: A flap partially covering the end in “iformes” (for example, Passeri- communally, with several females lay- nares; it may help to keep out debris. formes). [1·52] ing eggs in the same nest. [1·83]

Cornell Laboratory of Ornithology Glossary Oti – Par 45 otic: Of or relating to the ear. [4·11] the hard-shelled egg. In most birds, only the the small intestine; the pancreas secretes outbreeding depression: The reduction left oviduct is functional. [4·131] digestive juices into the small intestine in the degree to which local populations oxidation: The process by which a sub- and produces hormones (insulin, glu- are adapted to their local environments stance is chemically combined with oxy- cagon, and somatostatin) that regulate as a result of individuals dispersing and gen; also called “burning.” When body carbohydrate metabolism and blood mating with individuals in other popula- cells combine oxygen with the products sugar levels. [4·75, 4·122] tions (outbreeding). The reduction occurs of food digestion, such as carbohydrates, Pangea: The single land mass that formed because outbreeding individuals bring the reaction releases energy and two 245 million years ago from all the exist- alleles adapted to one local environment waste products—water and carbon di- ing continents and persisted throughout into another, where they may be less ad- oxide. [4·86] the Triassic period. [1·68] vantageous. [9·66] oxpeckers: Two bluebird-sized species in Pantanal: An enormous expanse of grass- outer shell membrane: A loose, fibrous the starling family (Sturnidae) that climb lands extending across parts of Brazil, membrane that lines the inner surface upon large grazing mammals in the Af- Paraguay, and Bolivia that each year is of the avian eggshell; it sticks tightly to rican savannas, removing ticks, insects, transformed into a huge marsh by tor- the eggshell, helping to hold it together. and the scabs of skin wounds—a behav- rential seasonal rains. It hosts one of the [8·62, 8·69] ior that benefits both the birds and their greatest concentrations of wildlife on the outer vane: The vane located on the side hosts. Also called tickbirds. [1·87] continent of South America, including of a wing or tail feather that is away from oxygenated blood: Blood whose red numerous waders and other water birds, the midline of the bird. On a wing feath- blood cells contain high levels of oxygen. as well as the few remaining Hyacinth er, the outer vane is on the edge of the Also called oxygen-rich blood, it is found Macaws. [1·73] wing that leads in flight. In flying birds, in all arteries except the pulmonary ar- pantropical: Distributed throughout the the outer vane is narrower than the inner tery and its branches. [4·81] tropics of the world. [1·81] vane, producing an asymmetry that aids oxygen-poor blood: See deoxygenated papilla (plural, papillae): A small bump. in flight. [3·3] blood. [4·81] 1. The bump at the end of each of a outer wing: The portion of the wing from oxygen-rich blood: See oxygenated bird’s deferent ducts where it opens into the wrist to the wing tip; the primary feath- blood. [4·81] the cloaca; also called the papilla of ers are located on the outer wing. [5·23] the deferent duct. When the cloaca is ova (singular, ovum): Female reproductive everted during copulation, the papillae cells, also called egg cells or eggs, both may slightly enter the female’s oviduct. before and just after they are fertilized by P [4·128] 2. One of many small bumps a sperm cell. [4·128, 4·130] Palaeognathae: One of the two Super- covering the surface of the skin of birds orders of ornithurine birds; it contains all during embryonic development. Each oval: Describes an egg that is shaped like birds with a palaeognathous palate—one papilla consists of a core of dermis and a chicken egg; also called ovate. [8·73] formed by large, rigid bones (compared a covering of epidermis and will even- oval window: Former name for the ves- to a neognathous palate in which the tually form an embryonic feather. Also tibular window. [4·57] vomer and basipterygoid process are re- called a feather papilla. [3·26] 3. One ovary: The female gonad; it matures duced and a flexible joint exists between of many spiny-tipped bumps on the un- and releases egg cells (ova) periodically the pterygoid and palatine bones). Palae- dersides of the feet of some birds, such as throughout the breeding season in a pro- ognathae includes all living ratites as well Osprey, to aid them in grasping slippery cess called ovulation, and also produces as the extinct elephantbirds and moas. prey. [3·45] 4. One of many tiny bumps the sex hormones estrogen, progesterone, [E·21, E·23] in the lining of the shell gland of the ovi- and testosterone. In most birds only the left palaeognathous: See Palaeognathae. duct; they secrete albumen and the hard, ovary is functional, and it enlarges greatly [E·21, E·23] calcium-rich outer shell around the fertil- during the breeding season. [4·128] ized ovum. [4·131] palate: The roof of the mouth. [4·91, ovate: See oval. [8·73] 4·112] parabronchi: Tiny (microscopic) air tubes formed by the branching of secondary Ovenbird: A wood-warbler (family Paru- palatine: Bony rod on each side of the bronchi within the avian lung; they form lidae) that breeds in mature deciduous upper jaw above the jugal arch (another a network within the lung tissue. Air car- forests of the eastern and central United set of bony rods). In birds, as the lower rying oxygen passes out of openings in the States and migrates to the Neotropics for jaw opens it moves the quadrate, which thick walls of the parabronchi and into a the winter. [8·35] pushes the palatine and jugal arch for- network of air spaces surrounded by a ovenbirds: A diverse Neotropical sub- ward, pushing on the premaxillary bones network of blood capillaries. Here, gas oscine family (Furnariidae, 240 species) to raise the upper jaw. [4·12] exchange occurs—oxygen dissolves into whose members are especially numer- Palearctic region: Zoogeographic region the blood and carbon dioxide and water ous in temperate South America; oven- that consists of most of the large landmass vapor move from the blood into the air and birds are named for the oven-shaped, of Eurasia, as well as northern Africa and back into the parabronchi, eventually to clay nests built by some species. [1·79] most of the Sahara Desert. In some sys- be exhaled. The avian system of airflow is overstory vegetation: The mature trees that tems, but not the one used by the Hand- much more efficient than that of mammals make up the canopy of a forest. [9·93] book of Bird Biology, Greenland is included (in which air exchange occurs in dead- oviduct: The tube that transports the egg in the Palearctic. [1·69, 1·70] end pockets called alveoli), because in from the ovary to (in birds) the cloaca; it is pamprodactyl feet: Foot arrangement birds air flows continuously across the suspended from the dorsal body wall by a in which all four toes, including the hal- surface of the capillary bed. [4·98] curtain-like membrane. After the egg enters lux, point forward. Found in some swifts. paramo: Humid grassland area of the the oviduct as an egg cell (ovum) and is fer- [1·21] high Andes in South America above the tilized, different sections of the ovary add pancreas: Digestive organ and endocrine tree line. It contains some shrubs and is different substances to the ovum to produce gland located in the uppermost loop of dotted with lakes and bogs. [1·75]

Handbook of Bird Biology 46 Par – Per Glossary parasitic: Describes a relationship be- species, all of which have a foot adapted such as owls, hawks, and kingfishers— tween two species or individuals in for perching on branches or stems. [1·66, and is regurgitated through the mouth; which one benefits as a result of some 7·25] also called a cast. [4·119, 4·120] cost to the other. [9·91] patagium: A fold of tough skin that extends pelvic girdle: The three bones (ilium, is- parasitoids: Animals (usually insects) from the brachium to the antebrachium; chium, and pubis) on each side of the that feed in or on a host animal for a long the patagium connects the shoulder to body that are partly fused with one an- period of time, consuming most of its the wrist and forms the leading edge of other and with the synsacrum in birds to tissues and eventually killing it. Some the inner wing in flight. [1·11] form a strong but lightweight attachment parasitoids (such as certain small wasps) patella: The kneecap; an ossification site for the muscles of the legs, tail, and live in the nests of cavity-nesting birds, within a tendon at the lower end of the abdomen. They also provide protection feeding on avian ectoparasites and actu- femur (thigh bone). The patella glides in for the abdominal organs. Also called the ally benefiting the birds. [8·58] a deep groove and adds stability to the pelvis. [4·24] parasympathetic system: The part of the knee joint. [4·24] pelvis: See pelvic girdle. [4·24] autonomic nervous system that acts on pecten: Highly vascularized structure of pendulous cup nest: A cup nest whose smooth muscle to reduce the heart rate the choroid layer of the eyes of all birds rim is supported but whose unsupported and to promote feeding, egg laying, and and some reptiles; it projects into the vit- belly (nest chamber) hangs from around 4 other “peaceful” activities, such as diges- reous body where the optic nerve exits inches (10 cm) to over 1 yard (1 m) below tion. Its nerves originate in the cranial the eyeball. The pecten is believed to the supports. It is generally entered from and sacral region. The other part of the nourish the retina and to control the pH the top. Pendulous cup nests are built by autonomic nervous system, the sympa- of the vitreous body. [4·47] New World orioles, oropendolas, and thetic system, functions under conditions caciques, as well as some Old World of stress. [4·35] pectinate claw: A modification of the side of the middle toe claw into a comb- weavers, and usually are woven from parathyroid glands: Several small endo- like, serrated edge thought to be used plant strips or fibers. Shallow pendulous crine glands located on or caudal to the as a preening tool. Found in only a few cup nests grade into pensile cup nests thyroid glands; they secrete parathor- birds, such as Barn Owls, nightjars, bit- (see separate entry). [8·32] mone, a protein that causes calcium re- terns, and herons. Also called a feather pensile cup nest: A cup nest whose rim is sorption from the bones. [4·74] comb. [3·46] supported but whose unsupported belly partial migration: Migratory pattern pectoral girdle: The three bones (clavi- (nest chamber) hangs below. Pensile cups in which some individuals in a popu- cle, coracoid, and scapula) on each side are built by New World blackbirds, vir- lation migrate while others remain as of the avian body that form a “free-float- eos, and kinglets; by many songbirds in year-round residents. When the specific ing” support for the wings. [4·19] Australia and Asia; and by a host of ori- individuals that migrate are determined oles and flycatchers and their relatives in genetically, a situation that occurs in en- pectoral veins: Paired veins (left and the Old World. Deep pensile cup nests vironments where the resources always right) that carry blood from the pectoral grade into pendulous cup nests (see sep- are sufficient to enable some, but not all, region; they merge with the subclavian arate entry). [8·30] and jugular veins on each side to form the individuals to overwinter, the migrat- perch gleaning: See gleaning. [6·44] ing birds are called obligate partial mi- cranial vena cavae leading to the heart. grants; examples include the European [4·83] perching birds: See passerines. [1·66, Robin and populations of the Blackcap pectoralis: Large, powerful flight muscle 7·25] in southern Europe. When the number of birds that attaches to the sternum. The perforate septum: A condition in which of birds the environment can support var- pectoralis has two portions: the larger the nasal septum (tissue separating the ies from year to year, facultative partial part pulls the wing down, slows it down left and right nasal cavities) has an open- migration evolves, in which the number at the end of the upstroke, and pulls the ing or is absent. In contrast with having an and identity of the individuals migrating wing forward; the smaller part pulls the imperforate septum (see separate entry), change from year to year in response wing down and back. [5·6, 5·7] having a perforate septum appears to in- to the availability of resources, usually peents: Short, nasal vocalizations given crease an animal’s sensitivity in detecting food. In this case, the individuals that from the ground at twilight by a male odors. However, it decreases the ability migrate are not determined genetically. American Woodcock as part of his court- to locate an odor’s source, because odors Facultative partial migration occurs in ship display. They are similar to the buzzy entering one nostril mix with those from the Blue Tit and some North American call of a Common Nighthawk. [7·16] the other. [4·90] chickadees. [5·56] pelagic: Of the ocean; pelagic birds pericardium: Thin membrane sur- partial molt: A type of molt in which only spend most of their life on the open sea, rounding the heart; its inner lining se- some of the feathers are replaced. [3·28] feeding at the surface or just below it cretes pericardial fluid, which reduces Passeri: One of the two large suborders and coming to land only to nest. Pelagic the friction of the beating heart against of Order Passeriformes (perching birds); birds include species in the order Procel- adjacent tissues. [4·80] members are also known as oscines, lariiformes, as well as tropicbirds, some perilymph: The fluid that fills the struc- songbirds, or true songbirds. Songbirds penguins, the boobies and gannets, most tures that make up the bony outer laby- have particularly complex voice boxes, alcids, the skuas and jaegers, the nod- rinth of the inner ear (the cochlea, semi- which allow them to sing more complex dies, kittiwakes, Sabine’s Gull, and some circular canals, and vestibule), in which songs than other birds. [7·25] terns. [1·98] floats the membranous labyrinth. [4·57] Passeriformes: See passerines. [1·66, pelagic feeders: See offshore feeders. period: A unit of geological time. Periods 7·25] [6·68] are divided into epochs, and successive passerines: All birds in the Order Pas- pellet: A compact ball of indigestible periods make up an era. [1·113] seriformes; also called perching birds. food, such as bones, fur, feathers, and in- peripheral nervous system (PNS): The The order contains approximately 4,600 sect exoskeletons, that is formed by the portion of the nervous system outside the species, nearly one-half the world’s bird gizzard of birds that eat meat or fish— brain and spinal cord; it consists of the

Cornell Laboratory of Ornithology Glossary Pes – Pop 47 cranial nerves as well as the spinal nerves pineal gland: Endocrine gland located in plexuses: Complex web- or net-like struc- and their associated ganglia. [4·40] the dorsal midbrain; it secretes the hor- tures along the spinal cord at the level of pessulus: Cartilaginous structure found mone melatonin, which plays a role in the front limbs or wings (brachial plexus) only in songbirds; it sticks up into the regulating daily activity cycles (circadian and hind limbs (lumbosacral plexus); syrinx at the bifurcation of the bronchi. rhythms). [4·71] they are formed by a merging of the large [4·93] pin feathers: Developing feathers that spinal nerves within each region. [4·39] phalanges (singular, phalanx): The bones are still surrounded by a feather sheath. plumage: 1. A bird’s entire feather coat. of the fingers and toes. [1·9, 4·25] [3·26] [3·33] 2. The set of feathers produced by a particular molt. With this usage, a bird phallus: See cloacal phallus. [4·129] pinyon-juniper woodland: Ecosystem of southwestern North America consisting of wears parts of two different plumages af- pharynx: Area, also known as the throat, park-like stands of small pinyon pines and ter a partial molt. [3·33] that begins at the back of the tongue and junipers growing on hills and mountain plunge diving: A foraging technique in connects the mouth and esophagus, and slopes below the montane forests. Five which an airborne bird dives under wa- where the digestive and respiratory path- species of breeding birds are particularly ter to pursue aquatic prey; performed by ways cross one another. The nasal cavities characteristic to this dry ecosystem: Gray birds such as Brown Pelicans, auks, gan- and auditory tube open into the pharynx. Flycatcher, Pinyon Jay, Juniper Titmouse, nets, Osprey, and kingfishers. [6·45] [4·91, 4·112] Bushtit, and Bewick’s Wren. [9·121] pneumatic: Describes bones that are Phasianidae: The pheasant family; it in- pipped: Describes an egg about to hatch, filled with air spaces and may contain cludes pheasants, partridges, ptarmigan, in which the embryo within has punc- air sacs; nearly all the bones of most birds grouse, guineafowl, turkeys, and Old tured a small hole. [8·105] are pneumatic. [4·5] World quail—as well as the Red Jungle- fowl (the wild relative of domestic chick- piracy: A foraging technique in which a pneumatic foramen: Opening in the avi- ens) and the spectacular peafowl. Its 177 bird steals food from another bird; used an humerus (upper wing bone) leading species are found throughout most of the by Parasitic Jaegers, frigatebirds, and to the air space within the bone; exten- world, but few species inhabit the Neo- some gulls, among others. [6·48] sions of the clavicular air sac lead into tropics. [1·89, 1·90] pitch: See frequency. [7·4] the pneumatic foramen and thus connect with the air space. [4·100] philopatry: The tendency of individuals pittas: An Old World family (Pittidae, 31 of certain species to eventually take species) of secretive, stocky suboscines PNS: See peripheral nervous system. up residence (and breed) near the area with long legs and a short tail; they live [4·40] where they were born. [9·65] on the tropical forest floor, and many podotheca: The tough skin covering the photoperiod: The amount of time during are brightly colored below and cryptic tarsus. [1·20] each 24-hour period that an organism is above. Pittas use their heavy bills to catch polarized light: Light in which the waves exposed to light. [5·63] a variety of insects and other small ani- are all oriented in the same plane, or in a mals, especially snails. [1·90] photoperiodism: A behavioral or physi- reduced number of planes, compared to ological response to day length. [4·140] pituitary gland: An endocrine gland at- unpolarized light. [5·94] tached to the ventral surface of the hypo- polarizing lens or filter: Something that phylogeny: The evolutionary history of thalamus; it consists of an anterior and a an organism. [1·32] selectively transmits only those light posterior lobe. The pituitary synthesizes, waves with certain orientations, creating phylum (plural, phyla): Level of classifi- stores, and releases a wide variety of hor- a polarized beam of light. [5·94] cation of organisms above “class” and mones. [4·72] polyandry: Mating system in which one below “kingdom”; similar classes are plankton: Microscopic plants (and al- placed within the same phylum. All birds female mates with several males within gae) termed phytoplankton and micro- the same breeding season. [6·77] are in the phylum Chordata. [1·52] scopic animals (and protozoa) termed physiographic regions: Regions of the zooplankton that float freely in aquatic polygamy: General term for a mating sys- world that are biologically distinct based environments. [1·101] tem in which individuals of one sex (ei- on climate conditions, topography, soil ther the males or females) mate with more plasma: The fluid portion of the blood; than one partner during the same breed- types, and plant communities; also called it contains sugars, inorganic salts, and ecoregions. [10·86, 10·87] ing season. Both polygyny and polyandry certain proteins (plasma proteins) found are forms of polygamy. [6·79] phytoplankton: Microscopic plants and only in the plasma, and carries numerous algae that float freely in aquatic envi- types of dissolved substances. Avian polygyny: Mating system in which one ronments; phytoplankton make up the plasma has a higher sugar, fat, and uric male mates with several females within “plant” portion of plankton. [1·103] acid content than the plasma of most the same breeding season. Polyandrous mammals. [4·86] species include Spotted Sandpipers, most pia: The innermost of the three vascu- jacanas, and certain phalaropes. [6·68] larized membranes called meninges platform nest: Nest consisting of a very that surround the brain and spinal cord. shallow depression in the top of a mound polymorphism: A situation in which one The meninges provide sustenance and of nest material. Varies in complexity species contains two or more distinct waste removal for the cells of the brain from the flimsy platform of twigs built by morphological types (morphs) of individ- and spinal cord, which are not served by Mourning Doves to the enormous stick uals, which are determined genetically. the circulatory system. [4·36] piles amassed by storks or large birds of Morphs may differ in physical charac- teristics such as color, body size, or bill pigeon’s milk: See crop milk. [4·116] prey such as Osprey and eagles. Some platform nests, such as those of some length or shape, but not in characteristics pigments: Colored substances that give terns and rails, float. [8·28] that are related to sex, age, locality, or color to the structures, such as skin and season. [9·42] feathers, in which they occur. In theory, playback experiments: Experiments in which researchers play songs (through a population: All the individuals of a spe- pigments can be extracted from these cies that live in the same area. [1·45, structures. [3·50] speaker) to territorial birds, and then note their responses. [7·48] 9·3]

Handbook of Bird Biology 48 Pop – Pro Glossary population density: The number of indi- With birds, technically used only within tervals that mimic the natural fire cycle of viduals per unit area in a given popula- the eye and inner ear. But in practice, often the landscape, to maintain certain types tion. [9·58] used interchangeably with caudal. [1·4] of habitats and the species that depend population dynamics: 1. Changes in popu- posterior air sacs: General term referring on them. [10·83] lation size, density, dispersion, and struc- to the air sacs nearest the bird’s caudal primary bronchus: See mesobronchus. ture over time. [9·49] 2. The study of how end—the posterior thoracic and abdomi- [4·98] populations change over time. [9·49] nal air sacs. [4·102] primary consumers: Level in a food chain population ecology: The study of how posterior chamber: Space between the or web that consists of organisms—usu- animal populations are related to, and re- iris and lens of the eye; it is filled with ally animals—that directly eat plants spond to, their environments. It involves aqueous fluid, which nourishes the eye or other producers; most primary con- monitoring and studying reproductive and removes wastes. [4·48] sumers are herbivores. [9·123] rates, survival rates, movements of indi- posterior lobe of the pituitary gland: Part primary feathers: The flight feathers of viduals and populations, and changes in of the pituitary gland that does not manu- the outer wing; they are attached to the population densities over time and from facture hormones itself, but instead stores manus. [1·11, 1·10] one area to another. [9·49] and releases neurohormones such as me- primary productivity: Gross primary population structure: The relative pro- sotocin and arginine vasotocin. [4·73] productivity is the rate at which green portions of males and females, individu- post-hatch brood amalgamation: A broad plants and the few other producers (such als of different ages, and individuals with term describing a variety of methods by as green algae) in an ecosystem produce other definable differences in a popu- which young from several broods may biomass, including the biomass that they lation. [9·49] combine into larger groups. [8·128] use themselves during respiration. Net population viability analysis (PVA): A postpatagium: A tough band of tendinous primary productivity is the gross primary computer technique developed by con- tissue running along the edge of the ante- productivity minus the biomass that the servation biologists in the 1980s that brachium and manus that trails in flight. producers use themselves; in other words, simulates the growth of populations over The postpatagium holds the primary and it is the rate at which the producers create time. Given numerous demographic vari- secondary feathers firmly in place and biomass that is available as food for other ables, such as birth and death rates at vari- supports each quill. [1·11] organisms. [9·87] ous ages and the annual variation in these powder downs: Down feathers that are primary succession: Ecological succes- rates, PVA can predict the probability that never molted but grow continuously, sion that begins on a substrate, such as local populations of different sizes will go disintegrating at their tips to produce a rock, sand, or lava, that has never before extinct over a designated period of time. fine powder resembling talcum powder, supported a community. Also refers to the The simplest PVAs are single-population which may help to waterproof the feath- process by which a lake gradually fills in models, which focus on just one popu- ers. [3·18] to form a bog community. See ecological lation (of different starting sizes and over succession. [9·110] different time periods) and ignore any powered flight: Type of flight in which dispersal of individuals between popu- the flying object produces thrust. In a primitive streak: In avian biology, very lations. More complex PVAs focus at the bird, refers to flight powered by flapping, early embryonic stage consisting of a metapopulation level (see separate entry); which produces thrust to propel the bird long trough (the “streak”) with raised these use more detailed information, such forward. [5·36] sides that establishes the cranial-caudal, as dispersal rates and patterns of behavior precocial: Describes young birds that left-right, and dorsal-ventral body axes. and mortality while moving across inhos- hatch in a relatively developed state— The primitive streak is formed in the flat- pitable habitats, to more realistically pre- downy and with their eyes open. Many tened mass of cells that lies on the upper dict the probability of extinction. [10·77] are soon able to walk or swim and even surface of the yolk. [8·64] eat on their own. [8·106] porphyrins: Pigments producing red, producers: Level in a food chain or web brown, green, or pink colors in a num- precopulatory displays: Displays given that consists of green plants and a few ber of avian orders; they are especially by either sex immediately before copu- other organisms (such as green algae) that well known for producing the bright reds lation that appear to invite or solicit capture energy from the sun and then in- and greens of the turacos. Porphyrins are copulation. Female birds of many spe- troduce the energy into the food chain by complex, nitrogen-containing molecules cies perform a stereotyped precopulatory incorporating it into organic compounds, related to hemoglobin, that birds and display in which they raise the tail and which the producers synthesize from wa- other organisms synthesize by modify- hold the wings down or out, quivering ter and inorganic substances through the ing amino acids. [3·51] them. [6·40, 7·71] process of photosynthesis. [9·123] Porro prism binoculars: Binoculars in preen gland: See oil gland. [3·20] productivity: In biology, the total num- ber of grams of living material produced which the eyepieces are closer together preening: Feather maintenance behavior 2 than are the objective lenses. [2·38] in which a bird grasps a feather near its per square meter per year (g/m /yr) in a base, then nibbles along the shaft toward particular area or habitat. Productivity portal system: A blood circulatory pat- includes the increase in size of all living tern involving portal veins. [4·85] the tip with a quivering motion; this cleans and smooths the feather. Many things, as well as new individuals added portal veins: Veins (such as the hepatic birds gather on their bill oily secretions through birth or germination. Productivity portal vein and the renal portal vein) that from the oil gland, and then spread them may be measured as gross productivity carry blood between two capillary beds, on their feathers as they preen. [3·18] or net productivity (see similar definitions rather than connecting a capillary bed under primary productivity). [9·92] and the heart, as do most veins. Portal premaxilla: Main bone forming each side of the upper beak of a bird. The two projectile vomiting: Vomiting that is veins allow a second organ to process sufficiently forceful to propel the regur- the blood before it is returned to the heart halves together are called the premaxil- lary bones. [4·11] gitated substance a short distance away. and thus the general circulation. [4·85] Birds such as vultures, herons, gulls, and posterior: Toward the back of an organ- prescribed burning: Purposefully starting tubenoses (Fulmars, albatrosses, petrels, ism, using the earth as a frame of reference. fires under controlled conditions, at in- storm-petrels, and diving petrels) use this Cornell Laboratory of Ornithology Glossary Pro – Rec 49 technique to ward off predators; Fulmars tures convergent with those of birds, such vertebrae. The pygostyle is shaped like a can spew an oily, foul-smelling mixture as hollow bones and a slight keel on the plowshare and provides an attachment site of flesh and fluid several feet. [6·54] sternum, but their large, membranous for the flight feathers of the tail. [4·18] proprioception: A “body-parts-position wings were structurally unique: they pylorus: A muscular sphincter (circular sense”: the subconscious perception of were supported by a single, greatly elon- band of muscle) at the lower end of the impulses from the muscles, tendons, and gated fourth finger and attached to the stomach; it regulates the passage of food joints that allows the body to know the side of the body and possibly also to the from the stomach into the small intestine. position of its parts, which is crucial for hind limb. [E·33] [4·120] balance and movement. [4·33] pterylae (singular, pteryla): Areas of a pyriform: Pear-shaped. Describes an egg proteins: Complex molecules composed bird’s skin where feathers are attached; that is rounded at one end and relatively of strings of amino acids; proteins are the also called feather tracts. [3·2] pointed at the other; also called conical. main building blocks of all living organ- pterylosis: The arrangement of feather [8·73] isms and also act as enzymes, assisting tracts and bare patches on a bird. [3·2] chemical reactions. [1·41] pubis: One of the three paired bones that protoplasm: The living material compos- fuse to form the pelvic girdle; the long, ing all organisms. [4·144] thin pubis runs caudally along the ven- Q quadrate bone: Bone on each side of the tral side of the ischium, and is only partly proventriculus: The upper part of the bird’s skull, at the caudal and lower end of the fused with it. [4·24] two-part stomach; it is elongate and glan- upper jaw; in birds, it links with the ar- dular, and secretes mucus, hydrochloric puddle ducks: See dabbling ducks. ticular bone on each side of the lower acid, and an inactive precursor to pepsin, [6·47] jaw. This linkage forms the joint between an enzyme that digests protein. [4·118] pulmonary arteries: Two large arteries the upper and lower jaws, allowing a bird proximal: Toward the center of the body (right and left) carrying oxygen-poor to open its mouth widely. [4·12] (the elbow is proximal to the fingers) or blood from the pulmonary trunk (com- quill: Feather. [1·11] toward the origin of a structure (the base ing from the right ventricle of the heart) of a feather—where it is attached—is to the lungs; the pulmonary arteries and proximal to its tip). [1·4] their branches are the only arteries that proximate: In biology, pertaining to im- carry deoxygenated blood. [4·78] R mediate, internal causes; mechanistic. pulmonary arterioles: Tiny arteries with- race: A population of a species, usually Proximate questions about a behavior, in the lung that supply deoxygenated in a particular geographic area, that is for example, look into the internal causes blood to the capillaries surrounding the morphologically distinct from other pop- of the behavior—how it is carried out. For network of air spaces adjacent to the ulations of the same species, but whose comparison, see ultimate. [6·2] parabronchi, where air exchange occurs members remain capable of interbreed- in birds. [4·99] ing with members of those other popula- proximate factors: In biology, the actual tions. Also called a subspecies. [1·55] cues that trigger the body to carry out a pulmonary circulation: Portion of the cir- particular process or behavior. For exam- culatory system carrying blood from the rachis: The portion of the feather shaft ple, day length is a proximate factor that heart to the lungs and back to the heart. above the calamus; it is solid and has a affects the timing of breeding in birds. For [4·77] vane on each side. [3·3] comparison, see ultimate factors. [8·11] pulmonary trunk: A large artery that car- radiale: The larger of the two wrist bones Prunellidae: A family of 13 sparrow- ries oxygen-poor blood from the right of birds; it consists of fused carpals. like birds with slender, pointed bills; it ventricle of the heart and then branches [1·9] includes the accentors and Dunnock. to the right and left lungs as the right and radius: The smaller of the two bones Prunellidae is the only bird family en- left pulmonary arteries. [4·78] supporting the antebrachium (forearm). demic to the Palearctic region. [1·70] pulmonary veins: Two large veins (left [1·9] pruning: A foraging technique in which a and right) carrying oxygen-rich blood range: The geographic area within which bird bites off and eats plant buds; used by from the lungs to the left atrium of the a species or population generally re- Ruffed Grouse and other birds. [6·46] heart; the only veins (along with the pul- mains at a particular time of the year; a pseudo-babblers: A family (Pomatos- monary venules) that carry oxygenated species may have different breeding and tomidae, 5 species) of noisy, busy, social blood. [4·78] nonbreeding ranges. Also called the geo- ground-feeding songbirds with long, pulmonary venules: Tiny veins within graphic range. [2·19, 9·49] curved beaks and long, towhee-like tails; the lung that carry oxygenated blood raptors: Members of the orders Falconi- they are endemic to the Australasian re- away from the capillaries surrounding formes and Strigiformes, which contain gion. [1·95] the network of air spaces adjacent to the all the diurnal and nocturnal birds of pseudosuchian thecodont: Term used pri- parabronchi, where air exchange occurs. prey. [1·66] marily by Robert Broom in his “pseudosu- [4·99] ratites: All birds lacking a keel on the chian thecodont hypothesis” to refer to the pupil: The opening in the center of the sternum. Includes flightless birds—the thecodonts (see separate entry). [E·33] iris; it controls the amount of light enter- Ostrich, rheas, Emu, cassowaries, and pseudosuchian thecodont hypothesis (of ing the eye. [4·47] kiwis—as well as the tinamous, which bird evolution): Theory that birds evolved pus: The yellowish fluid seen in infec- are fully capable of flight. [E·22] approximately 230 million years ago from tions; it is a mixture of bacteria, dead rattle: A harsh trill. [2·16] small, arboreal thecodonts; first proposed white blood cells, and fluid. [4·88] recolonization: The establishment of a by Robert Broom in 1913. [E·8] PVA: See population viability analysis. population of a particular species in an pterosaurs: Flying reptiles from the Tri- [10·77] area where that species formerly oc- assic that radiated and diversified in the pygostyle: The tail bone of birds; it is curred, generally through natural dis- Jurassic and Cretaceous. They had fea- formed by the fusion of the final few caudal persal. [10·77]

Handbook of Bird Biology 50 Rec – Rol Glossary rectrices (singular, rectrix): The long, stiff releasers: Specific objects, physical reverse sexual size dimorphism: A situ- flight feathers of the tail. [1·12, 1·13] features, or behaviors that activate (or ation in which females are significantly recurrent bronchi: Secondary bronchi “release”) a specific response in an in- larger than males of the same species; (branches off the main air tube, the me- dividual. [6·5] found in many predatory birds such as sobronchus) within the avian lung that relict population: A small, isolated popu- raptors, jaegers, skuas, frigatebirds, and connect to the air sacs outside the lung. lation that is a remnant of a much larger boobies. Because the sexes are different [4·98] population that once existed over a wider sizes, they can take different sizes and geographic area. [9·57] thus different types of prey. This may re- recurved: Curved upward. Describes duce competition between mates, but beaks such as that of the American Avo- remiges (singular, remex): The longest why females of these species are larger cet. [1·17] wing feathers, also called the flight feath- than males (rather than the more typical red blood cells: Flattened, elliptical, red- ers of the wings; they include the primary situation, in which females are smaller) colored cells found in the blood; they and secondary feathers. [1·11, 1·13] remains a matter of much speculation. house the iron-containing pigment he- renal efferent veins: Veins that collect [9·40] moglobin, which carries large amounts blood from the capillary beds of the kid- rhamphotheca: Outer covering (sheath) of oxygen and gives the blood its red neys and carry it to the caudal vena cava, of the beak; it grows throughout a bird’s color. The red blood cells of birds have a which returns it to the heart. [4·85] life. In most birds it is hard and horny, but nucleus, unlike those of mammals. Also renal portal system: System of blood in waterfowl, pigeons, and some shore- called erythrocytes. [4·86] flow, found in birds and reptiles but not birds it is soft and leathery. [3·39] red fibers: A type of muscle fiber that in mammals, that collects venous blood rheas: Two species (family Rheidae) of appears red because it contains a large from the lower portion of the digestive large, flightless ratites of temperate open amount of myoglobin (which carries oxy- tract via the caudal mesenteric vein, and country in South America. [1·74] gen) and is permeated by massive cap- conveys it to the venous ring at the kid- illary beds (containing many red blood neys, where some of it passes through rhomboid sinus: Opening on the dorsal cells, which also carry oxygen). Muscles a capillary bed before being conveyed midline of the lumbosacral enlargement with many red fibers produce energy aer- back to the heart. The reason for this pat- of the spinal cord; it contains a gelatinous obically (using oxygen), without building tern of blood flow is unknown. [4·85] mass of neuroglial cells rich in glycogen, up the quantities of lactic acid that cause known as the glycogen body, whose renal portal veins: Two veins (left and function is unknown. The rhomboid si- fatigue, and thus can sustain actions for right) carrying blood from the caudal long periods of time. Therefore, long-dis- nus and glycogen body are unique to mesenteric vein to the capillary beds of birds. [4·39] tance fliers often have a preponderance the two kidneys; part of the renal portal of red fibers. Muscles with many red fi- system of birds. [4·83] rictal bristles: Stiff, hairlike feathers pro- bers often are called “dark meat” (as in jecting from the base of the beak in birds the drumstick of a turkey). [5·7] repertoire: See song repertoire [7·14, that catch insects. They may protect the 7·19, 7·85] or vocal repertoire. [7·10] redirected activities: Actions that are face and eyes of some birds that capture appropriate for a given situation, but are residents: Individuals that live year round large, scaly insects; they also may help directed at an inappropriate subject; often in a particular area. [5·54] birds detect movements of prey held in they appear at times of stress or conflicting resource-defense polygyny: Mating sys- the beak. [3·13] motivations. Some have been exaggerated tem in which one male may have several right heart: The right atrium and right and incorporated into displays. [6·33] mates, and the females choose their male ventricle of the heart; these chambers reflex: A relatively simple and stereo- partners by evaluating the quality of their contain deoxygenated blood, which typed response to a stimulus; it may be breeding territories; males with better ter- does not mix with the oxygenated blood either automatic or learned. [4·34] ritories attract more mates. This system of the left heart (left atrium and ventricle). may evolve when safe nesting sites or rich [4·78] refugia: Areas remaining stable when sur- food supplies are more important to the ritualization: The evolutionary process rounding areas are undergoing change survival of the young than is male paren- (such as that caused by glaciation); often by which certain everyday motions be- tal care. Found in Red-winged Blackbirds come exaggerated, repeated, and ste- used to describe isolated patches of rain and a few other species. [6·73] forest separated by grasslands during gla- reotyped into displays presenting a clear cial periods. [1·77] reticulate podotheca: A podotheca that message. [6·32] is divided into a network of small, irregu- rockfowl: Dull-colored African song- regional population: A population that lar, nonoverlapping plates. [1·20] inhabits a fairly large area; for example, birds (family Picathartidae, 2 species) all the Canada Geese in the central mi- retina: The innermost layer of the eyeball, with colorful, bare heads; they hop along gratory flyway. [9·49] upon which images are focused; it is pig- the rain forest floor in rocky areas and mented and contains the light-sensitive nest colonially in caves. [1·85] reintroduction: The establishment of rod cells and cone cells. [4·49] individuals of a species (through human rods: One of the two kinds of light-sensi- effort) in an area where that species used retort nest: A globular nest with an en- tive cells lining the retina of the eye; they to live—using individuals from a different trance tunnel; built out of mud by many are particularly good at detecting low area (translocation) or from a captive mud-nesting swallows such as Cliff Swal- light levels (but not in differentiating col- breeding program. [10·89] lows, and constructed from grass by many ors, which is the function of the cones). African weavers and New World swifts in When a rod cell is stimulated by light en- relative abundance (of species): The the genus Panyptila. [8·36] ergy, it sends a nerve impulse to the brain number of individuals of each species via the optic tract. [4·48] living in a particular community or area reverse Porro prism binoculars: Binoc- compared to the numbers of individuals of ulars in which the objective lenses are rollers: Old World nonpasserines (family other species in that area. Relative abun- closer together than the eyepieces; char- Coraciidae, 12 species) colored in shades dance measures how evenly individuals acteristic of some compact binoculars. of blue, pink, olive, or chestnut; they are are distributed among species. [9·82] [2·38] named for their rolling or rocking dives during courtship flights. They have long, Cornell Laboratory of Ornithology Glossary Roo – Sea 51 slender beaks and forage by perching and chambers contain hair cells and dense composed of two groups, the herbivorous then flying to the ground to catch large crystals called statoconia, both of which sauropods and the carnivorous theropods. arthropods. [1·87] are embedded in a gelatinous material [E·8, E·33] roof prism binoculars: Binoculars with that is surrounded by endolymph. The Sauriurae: One of the two major sub- straight barrels, in which the eyepieces statoconia and hair cells perceive the po- classes of birds; it contains primitive birds and objective lenses are directly in line. sition of the head with respect to gravity such as Archaeopteryx and the enantior- [2·38] (see entry for statoconia). [4·58] nithines (opposite birds), but no living rostral: Toward the beak; used for posi- sacral vertebrae: The vertebrae of the bird groups. [E·20] tions on the head and neck. [1·4] pelvic region; in birds, they are all fused sauriurine: Describes birds in the Sub- with the lumbar vertebrae and with some class Sauriurae. [E·20] round window: Former name of the co- of the thoracic and caudal vertebrae to chlear window. [4·59] form the synsacrum. [4·15, 4·18] scapula: The shoulder blade; one of the r-selected species: Species in which in- three sets of paired bones that make up sagebrush ecosystem: Fairly dry ecosys- the pectoral girdle. [4·19] dividuals have a life history strategy that tem found above the lowest elevations emphasizes high rates of reproduction of the Great Basin of North America; it scapulars: A group of feathers in the instead of (and probably at the expense is a shrubland dominated by sagebrush, shoulder region. [1·12, 1·13] of) long-term survival. These species are shadscale, and other woody growth 2 scavengers: Organisms that eat dead or- generally small and tend to develop rap- to 5 feet (0.6 to 1.5 m) high. Of the few ganisms and wastes. [6·63] idly, breed at an early age, have many breeding birds, four species are particu- scientific method: A procedure that sci- young per cycle (large litters or clutches), larly characteristic: Sage Grouse, Sage breed frequently, care for their young entists use to investigate how the world Thrasher, Sage Sparrow, and Brewer’s works. The specific steps typically fol- only briefly, and have short life spans. Sparrow. [9·121] [9·45] lowed by scientists to investigate aspects sagittal plane: A vertical plane through of the world vary among the different sci- rump: The lower back of the bird, above the long axis of an organism, extending entific disciplines, but in the biological the tail coverts; conspicuously colored in from head to tail. It divides the body into sciences the scientific method usually some birds. [1·6] left and right portions. [1·4] involves the following: asking a question runaway selection: One possible expla- salivary glands: Glands that secrete sa- about the world, formulating the ques- nation for why females of some species liva, which primarily acts to moisten food tion into a testable hypothesis, designing choose the males with the most elaborate in the mouth. Some birds that obtain their a study and collecting unbiased data, an- ornaments (such as ornate plumage) to food from the water, such as Anhingas, alyzing the data to see if the hypothesis is copulate with. The explanation applies have no salivary glands. The salivary supported or rejected (this often involves primarily to females choosing nonpater- glands of others, such as certain swifts statistical testing—see separate entry), nal sexual partners, either (1) males for ex- and swiftlets, secrete an adhesive sub- and then drawing conclusions. [6·97] trapair copulations, or (2) mates in species stance used in nest building. [4·111] sclera: Tough, whitish, outer layer of con- in which males do not provide parental sally gleaning: See gleaning. [6·44] nective tissue surrounding the eyeball. care or other resources for their offspring [4·48] (such as territories with food or nesting sandgrouse: Sixteen bird species that sites). The runaway selection hypothesis form the family Pteroclidae and that scleral ossicles: Bony rings supporting suggests that females may choose males inhabit the deserts and dry savannas of the eyes of all birds, lizards, turtles, and with elaborate ornaments simply because Africa and Eurasia; they are similar in fishes; they are not present in mammals. their sons will then inherit these attributes size and shape to pigeons, but taxono- [4·47] and be especially attractive to females, mists are not sure of their relationships to scrape: A shallow depression created by and thus produce more offspring (of both other families, and thus their order is not certain types of birds for use as a nest. It sexes); in this way, both the genes for pos- determined. Sandgrouse feed primarily usually contains little or no nest material; sessing elaborate ornaments and the genes on dry seeds and often fly long distances in some species it may be lined with a few for choosing male partners with those to water holes to obtain water. They are flat pebbles. Scrapes are built by many ornaments will increase in frequency in well known for their specialized belly plovers, terns, skimmers, and penguins. future generations. As each generation of feathers, which are structurally adapted [8·26] females chooses the most ornate males to absorb and retain water. Parents (usu- screamers: A South American family available, the ornaments evolve to be ally the male) soak the belly feathers at a (Anhimidae, 3 species) of heavy-bodied, more and more elaborate, and selection water hole, and then carry the water back goose-like birds with far-reaching calls. is said to “run away” with the trait—even to their nestlings. [8·133, 9·21] [1·74, 4·22] to the extent that having an elaborate or- sap well: A rectangular hole up to a few scrub desert: Hot, dry ecosystem of the nament may reduce a male’s longevity. For inches long, drilled through the bark of lowlands and valley floors of southwest- runaway selection to get started, females trees and shrubs by a sapsucker. The bird ern North America; it is composed of must gain some other advantage by choos- may return to the well many times to bare ground with widely spaced plants ing males with elaborate ornaments than feed, both on the sap that flows from the such as creosote bush, ocotillo, and bur simply having attractive sons. Once the wound and on insects that are attracted yucca. The scrub desert ecosystem hosts trend is begun, however, the advantage to the flowing sap. Other birds and mam- a moderate number of breeding birds, need no longer exist for runaway selection mals also may feast at the sapsucker’s which tend to synchronize their nesting to proceed. [6·84] well. [9·128] with periods of rainfall. [9·122] saurischian dinosaurs: One of the two scutellate podotheca: A podotheca that major groups of dinosaurs, the sau- is broken up into overlapping scales. rischian dinosaurs are also known as [1·20] S “reptile-hipped” dinosaurs because of sacculus: One of the two membranous seabirds: All birds directly associated the structure of their hip joint. They are chambers inside the bony vestibule of the with the open seas and consistently de- inner ear (the other is the utriculus); these Handbook of Bird Biology 52 Sea – Sif Glossary pendent on the seas for food; also called sense changes in the organism’s speed or sexual selection: Form of natural selec- marine birds. [1·65] direction, thus guiding balance. [4·58] tion that occurs when individuals differ search image: A general idea or internal semicircular ducts: Three ducts floating in their ability to acquire mates. In sexual representation of an object being sought in the perilymph of the semicircular ca- selection the characters that enhance that allows an animal to find it easily. If an nals of the inner ear, one in each plane mating success (either by increasing animal has a search image for a particular of space; they contain the hair cells that an individual’s ability to compete with type of prey, it is able to find the prey more guide balance by sensing changes in the other members of the same sex and thus efficiently than it can find other prey for organism’s speed or direction. They are gain mates, or by directly increasing an which it has no search image. [9·25] part of the membranous labyrinth of the individual’s attractiveness to the opposite sex) are selected for, in some cases even secondary bronchi: Branches off the me- inner ear and thus are filled with endo- lymph. [4·60] if they decrease individual survival (see sobronchus (main air tube) of each avian runaway selection). [3·68, 6·82] lung; some connect to the air sacs and are semilunar membrane: Membrane that called recurrent bronchi, and others divide extends from the pessulus into the cavity shaft: The stiff, central rod of a feather, to form the tiny parabronchi, the major re- of the syrinx; it is found only in songbirds. from which the vanes extend. [3·3] spiratory units of the avian lung. [4·98] [4·93] shell gland: Region of the avian oviduct secondary cavity nesters: See cavity semilunar valves: Valves, each com- after the isthmus and before the vagina; it adopters. [8·39] posed of three cusps (flaps), that are lo- is lined with papillae that secrete fluid al- cated at the beginning of the pulmonary bumen and the calcium-rich shell around secondary consumers: Level in a food the fertilized ovum. Shell pigments, if chain or web that consists of organ- trunk and the aorta; the semilunar valves prevent the backflow of blood into the any, also are added here; the patterns of isms—usually animals—that directly eat pigmentation reflecting the speed of the primary consumers. [9·123] ventricles as the ventricles relax after each heartbeat. [4·79] egg’s passage and rotation through the secondary feathers: The flight feathers of region. [4·131] semiplumes: Feathers that occur in a the inner wing; they are attached to the shivering: Uncoordinated muscle fiber ulna of the antebrachium. [1·11, 1·13] continuum of forms between down and contour feathers. Located at the edges of contraction; a way to produce heat by secondary sex characters: Anatomical the contour feather tracts, semiplumes muscle contraction without directed features that distinguish the sexes, but are provide insulation and help to maintain movement. [4·27] not directly involved in the production of the bird’s streamlined shape. [3·17] shock molt: A rare situation in which eggs or sperm. [4·137] sensitive period: A relatively brief span of a bird sheds many feathers all at once; secondary succession: Ecological suc- time early in life during which songbirds it usually is the result of stress, such as cession (see separate entry) that begins are best able to memorize the details of when a bird is grabbed by a predator (tail with bare soil or an existing community. songs they hear. [7·26] feathers may be shed if grabbed by the [9·110] tail), handled by humans, or exposed sensory neuron: Nerve cell that conveys to violent natural events such as earth- Secretary-bird: A long-legged bird of impulses to the spinal cord and brain; prey that stalks the African savannas on quakes and tornados. Also called fright these impulses are interpreted as sensa- molt. [3·38] foot for snakes, other small vertebrates, tions—either conscious or subconscious. and large insects. It is the only member of [4·33] Shoebill: A large, stork-like water bird its family, Sagittariidae. [1·84] endemic to Africa; it has a huge, shoe- sentinel system: A system in which group shaped, hooked bill for seizing large fish. Section 404, Clean Water Act: Part of members take turns watching for danger. the Clean Water Act (see separate entry) It is the only member of its family, Balae- Birds using sentinel systems include nicipitidae. [1·84] passed in the United States in 1972; Sec- Florida Scrub-Jays and some American tion 404 strives to protect wetlands by Crows. [6·56] shorebirds: Oystercatchers, plovers, regulating the filling of, or discharge of snipes, sandpipers, curlews, phalaropes, dredged material into, all fresh waters of septum: Tissue that extends ventrally and sheathbills. Ornithologists in Britain the United States, including wetlands. from the upper wall of the nasal cavities and the British Commonwealth, except It allows wetland filling by permit if and separates the left and right nasal cavi- Canada, speak of shorebirds as “waders.” applicants show that less damaging al- ties from one another; the septum may [1·65] ternatives are not feasible, and that the be perforate or imperforate (see separate entries). [4·90] shore feeders: Birds (or other animals) water will not be degraded significantly. that hunt on shore in the intertidal zone Applicants must avoid wetland impacts seriemas: A family (Cariamidae, 2 spe- (between the low and high tide marks), when possible, minimize wetland dam- cies) of fast, long-legged, grassland birds relatively close to their nests; shore feed- age, and provide compensation (wetland of South America; seriemas chase down ers include marine waders, such as the mitigation—see separate entry) for all their reptile prey on foot. [1·75] American Oystercatcher, and many other wetland impacts deemed unavoidable. sex hormones: Hormones produced by shorebirds. [6·68] [10·93] the anterior lobe of the pituitary gland, siblicide: The killing of one’s sibling; in seedsnipes: A family (Thinocoridae, 4 the adrenal glands, and (primarily) the birds whose offspring practice siblicide, species) of ptarmigan-like birds that nest gonads; they regulate the anatomical usually the stronger, older nestlings kill both in the southern lowlands and the structures, physiological processes, and the younger, weaker nestlings, either by high mountains of South America. [1·74] behaviors that are essential for repro- direct attack or by pushing them out of semicircular canals: Three bony, ring- duction. [4·137] the nest. [6·87] like canals of the inner ear, arranged at sex ratio: The ratio of males to females sifting: A foraging technique in which a right angles to one another (one in each (or females to males)—usually noted for bird sweeps its partly opened bill from plane of space); they are part of the bony a given population. [9·80] side to side in water or mud, straining out labyrinth, and thus are filled with peri- sexual dimorphism: A situation in which small animals and plant material with the lymph. They enclose the semicircular males and females of the same species dif- specialized edges of the bill as the wa- ducts, which contain the hair cells that fer from each other in size or form. [9·39] ter (or mud) drains out. Sifting is used by Cornell Laboratory of Ornithology Glossary Sin – Spe 53 Roseate Spoonbills, Northern Shovelers, smooth muscle: Type of muscle found intruders. Strictly speaking, songs are giv- and others. [6·46] in the walls of the hollow organs (such en only by songbirds—passerines in the single-population models (in PVA): See as the stomach or intestines) and in the suborder Passeri (oscines). In practice, population viability analysis. [10·77] walls of blood vessels larger than capil- however, similar vocalizations made by laries, especially arteries and arterioles. nonsongbirds (and even other animals) site fidelity: Loyalty shown by birds or The contraction of smooth muscle is not often are called songs. [7·14] other organisms to places they previously controlled consciously. [4·28] occupied; the places may be breeding lo- Southern Marine Region: The major fau- cations, nonbreeding locations, or stop- snood: The limp, red, fingerlike, fleshy nal region of the seas that includes the over points between the two. Also called structure that projects over the bill from frigid waters around Antarctica north to site tenacity. [5·75, 9·13] the forehead of a Wild Turkey. [3·48] about 35 degrees south latitude. [1·99, 1·100] site tenacity: See site fidelity. [5·75] soaring: In a bird or other animal, fly- ing without flapping the wings or limbs, southwestern oak woodland: North skeletal muscles: Muscles that move the while gaining altitude or remaining hori- American ecosystem that is scattered bones; they constitute the “meat” of an zontal (as compared to gliding, in which on hills and mountain slopes and that animal and are under conscious control. the animal loses altitude). [5·36] contains open woodlands consisting Also called voluntary muscles. [4·26] social foraging: See cooperative forag- primarily of oaks and occasional large skulling: A technique by which field ing. [6·64] ponderosa pines; southwestern oak workers use the degree of skull ossifica- woodland is found in Utah, Nevada, Cal- tion and pneumatization to determine Solnhofen limestone: Fine-grained lime- ifornia, New Mexico, Arizona, and parts the age of live passerines captured in the stone deposits near the village of Soln- of Colorado. Rainfall is low to moderate. fall. [4·10] hofen in Bavaria that, due to meticulous [9·120] mining, have produced an array of amaz- slope soaring: A type of static soaring in ingly well-preserved fossils of plants, in- spatial learning: Learning the location of which a bird derives lift from air deflected vertebrates, fishes, and reptiles, as well objects. [6·12] upward when wind strikes a hill, ridge, or as several specimens of Archaeopteryx. speaker replacement experiments: Ex- cliff; or from rising eddies created when [E·2] periments in which researchers remove wind spills over a cliff. Slope soaring is soma: See cell body. [4·32] animals (usually male birds) from their particularly common along seacoasts territories, and then replace them with among gulls, terns, fulmars, and gannets; somites: In avian biology, paired seg- tape recorders and loudspeakers broad- it also is common along mountain ridg- ments that appear along the middle casting various sounds (the male’s song, es, where ravens, crows, and migrating region of the cranial-caudal axis of the the song of a different species, back- hawks may soar. [5·42] very early embryo; eventually they will ground noise with no song, and so on), slots: Gaps between the feathers of the become vertebrae and muscle. [8·65] depending on the specific experiment. wing tips, created when a bird having sonagram: A visual representation of Then the researchers watch the reaction narrow-tipped primaries spreads them sounds, such as bird song, plotted as a of other birds in the area. [7·69] during flight. Slots are most common graph of frequency or pitch (vertical axis) speciation: The formation (evolution) of in large, soaring birds, such as vultures, versus time (horizontal axis). The dark- new species through natural selection. condors, eagles, and certain hawks, be- ness of the images gives a rough indica- [1·57] cause they allow these birds, which have tion of the relative loudness of the dif- broad, rounded wing tips, to increase ferent sounds. Also spelled sonogram. species: 1. Level of classification be- their lift-to-drag ratio. Slots reduce tip [7·7] low “genus”; similar species are placed vortex by turning each primary feather within the same genus. A species usu- songbirds: Members of Suborder Passeri, ally is defined as a group of potentially into an individual, narrow, pointed “wing which is one of the two large suborders tip”; they also increase lift because each interbreeding individuals that share dis- of Order Passeriformes (perching birds); tinctive characteristics and are unlikely separated primary feather acts as an in- songbirds also are known as oscines or dividual airfoil. [5·35] to breed with other such groups of in- true songbirds. Songbirds have particu- dividuals. Distinguishing which groups slotted high-lift wings: Broad wings with larly complex voice boxes, which allow of organisms constitute species is not al- a deep camber and very prominent slot- them to sing more complex songs than ways easy, however, and slightly different ting; the broadness reduces wing loading other birds. [7·25] criteria may be used by different scien- and allows low flight speeds, and the slot- song control centers: Groups of nerve tists. [1·53] 2. The two-word designation ting increases lift, allowing the birds to cells within the brains of songbirds that constituting the Latin or scientific name carry heavy prey. Birds such as vultures, are involved with both the hearing and of a “species” (see definition 1); the first eagles, condors, storks, and some owls production of songs; they are intercon- word, capitalized, is the “genus” and the and hawks have this type of wing; they nected with one another in a series of second word, lowercase, is the “species” use it for static soaring. [5·39] complex pathways. Individuals that sing name (see definition 3). Both words are small intestine: The longest part of the frequently or that have large song rep- underlined or printed in italics. [1·47] 3. digestive tract, running from the stomach ertoires have large song control centers. The second word in the two-word scien- to the large intestine. The final processes The centers shrink in the nonbreeding tific name of a species (see definition 2). of digestion take place inside the small season—a time when most birds sing It is always lowercase, and is underlined intestine, as proteins are broken down less often. [7·39] or printed in italics. [1·47] into amino acids, carbohydrates are song repertoire: All the songs (different species account: A set of written field converted to simple sugars, and fats are song types) sung by an individual bird; for notes on a particular species. [2·52] reduced to glycerol and fatty acids. The comparison, see vocal repertoire. [7·14, species-area relationship: Typical re- small intestine is longer in birds that feed 7·19, 7·85] on foliage or grain than in those that feed lationship between the size of an area on fruit or meat, reflecting the difficulty songs: Loud vocalizations, often deliv- and the number of species it supports: as of digesting the cellulose in plant mate- ered from an exposed perch, that are pre- area size increases, so does the number rial. [4·120] sumed to attract mates or repel territorial of species. This relationship holds (1) for Handbook of Bird Biology 54 Spe – Stu Glossary patches of the same type of habitat either curs when an organism is already very 1 in 20—for example, 1 in 100 (.01)—the in the same or different areas (the larger well adapted to current conditions, and hypothesis is accepted. In the biomedical the habitat patch, the greater the number it tends to keep the species at an optimal sciences, where the risk of making a mis- of species it supports), and (2) for areas middle point. [1·37] take may carry serious consequences, ac- of increasing size centered on the same stable age distribution: Property of some ceptable levels of significance often are point (as the boundaries of the area be- populations such that the relative propor- set much lower. [6·97] ing considered are expanded outward, tions of individuals of different ages are the statoacoustic nerve: See vestibuloco- the area contains more species—both same each year, when the same time of year chlear nerve. [4·42] because the size of a patch of one type of (such as the start of the breeding season) is habitat may increase, and because a larg- statoconia: Dense crystals of calcium compared. Note, however, that in a stable carbonate embedded, along with sensory er area may contain more different types age distribution these proportions still of habitats). The species-area relation- hair cells, in a gelatinous material inside change predictably throughout each year: the utriculus and sacculus of the inner ship also holds for islands surrounded by in most birds, for example, the proportion water. [9·95, 10·72] ear. As the head’s orientation changes of very young individuals increases in late with respect to gravity, the crystals settle species population: A population of all spring as eggs hatch, and then gradually against different hair cells, stimulating the living individuals of a particular spe- decreases as many juveniles die before them; the brain determines the head’s cies; also called simply a species. [9·49] reaching one year of age. [9·80] position by detecting which hair cells species richness: The number of species star compass: The mechanism by which are stimulated. [4·61] living in a community (or in a particular nocturnally migrating birds are able to stepping stones: In conservation biology, area). The more species, the greater the use the star patterns surrounding the small pieces of habitat, suitable for a par- species richness. [9·82] North Star (in the Northern Hemisphere) ticular species, that are located between sperm: See spermatozoa. [4·127] to determine which way is north (and thus larger habitat preserves; they allow indi- the other compass directions as well). A spermatozoa (singular, spermatozoan): viduals to move between the larger pre- young bird initially learns how to use the serves. [10·80] Sperm cells or sperm; each spermatozo- star patterns by observing their rotation an is a single cell composed of a DNA- and using the point at the center of their sternal rib: The ventral portion of each containing head and a propulsive tail. rotation (the North Star, Polaris) as the thoracic rib; it articulates with the ster- Sperm are short and simple in nonpas- direction to North; but once a bird learns num ventrally, and with the vertebral rib serines, and longer and spiral-shaped in the star patterns, it can find north even in dorsally. [4·18] passerines. [4·127] a stationary planetarium. [5·86] sternum: The breastbone; it provides an spinal cord: A cable of neurons con- statant cup nest: Cup nest constructed attachment site for the ribs and pectoral ducting nerve impulses to and from the on top of a hard physical support or sup- muscles. In all flying birds except tina- brain; it runs inside the vertebral canal ports, including the ground, building mous, the sternum has a keel (carina). of the vertebral column, extending from ledges, or tree branches; built by Amer- [4·23] the medulla oblongata of the brain to the ican Robins, Horned Larks, and many stochastic: In biology, describes events tail. [4·38] other birds. [8·30] whose occurrence and/or outcomes are spinal nerves: Nerves that arise from the static pressure: The force, produced by the result of random (or at least unpre- spinal cord in pairs—one on each side random motion of molecules, that air dictable) factors. For comparison, see of the cord—all along the spinal cord; exerts uniformly in all directions. When deterministic. [9·77, 10·67] each pair carries sensory input and motor you squeeze a balloon, you feel the static stomach: Saclike, expandable digestive output, and serves a very specific region pressure from the air inside. [5·11] organ between the esophagus and the of the skin, or very specific muscles or small intestine; in birds, the stomach other organs. [4·42] static soaring: A type of soaring (flying without flapping or losing altitude) in consists of two parts: the proventriculus spinous process: A ridge of bone project- which birds take advantage of the energy (upper) and gizzard (lower). [4·118] ing from the dorsal surface of each ver- in rising air masses (either in thermals, stooping: A foraging technique used by tebra; it is particularly well developed in called thermal soaring, or along hills or falcons in which a bird drops through the the thoracic vertebrae. [4·16] ridges, called slope soaring) to obtain lift air at great speed in pursuit of a flying bird spotting scope: A medium-range tele- with little or no energy expenditure on or insect. [6·45] scope commonly used for bird watch- their part. Performed by birds with slotted structural colors: Colors not produced ing, usually with magnification power high-lift wings. [5·39] by pigments, but instead by the reflection between 15x and 60x. [2·42] statistical testing: Mathematical pro- of certain wavelengths of light off the ac- spread-wing posture: A stance in which cedures used by scientists to determine tual physical structure of the object (such a bird stands motionless with the wings whether a set of observations or the results as a feather of a bird or a scale of a but- extended to the side, either to dry the of an experiment either (1) support their terfly wing). [3·50] wings or to absorb sunlight. Adopted by hypothesis, or (2) could result from chance studbook: A collection of information many large birds, such as cormorants, an- alone. Most statistical tests generate a on the individuals of a particular spe- hingas, pelicans, storks, and New World number that indicates the probability that cies. Key to a studbook is data on the vultures. [3·22] the observations or results could have relatedness among individuals—for spurs: Bony outgrowths from any part of occurred from chance alone. The lower example, which individuals are the par- the skeleton, such as the wing (see wing the probability, the greater the statistical ents, siblings, and offspring of a given spurs). [4·22] significance of the data. Many scientists individual. But studbooks may contain a choose a 1 in 20 probability (.05) as the square tail: Tail shape in which the rectri- variety of other information as well, such cut-off for significance: if the probability is ces are all about the same length. [1·19] as data on each individual’s disease his- greater than 1 in 20—for example, 2 in 20 tory, birth location and date, and location stabilizing selection: Natural selection (.10)—then the hypothesis is rejected, but in the wild or in captivity. Studbooks are that acts against any change in a current if the probability is equal to or smaller than used primarily by researchers who are characteristic. Stabilizing selection oc- Cornell Laboratory of Ornithology Glossary Sub – Syn 55 breeding endangered or declining spe- of the sun in the sky to indicate compass at different ages. The survivorship curve cies; the studbook information helps the direction. To do this, they must be able to shows the chance of mortality at different researchers to pair individuals with the compensate for the changing position of stages in an organism’s lifetime; the basic partners to whom they are least related, the sun during the day. [5·84] shape of the curve for a particular species thus promoting and preserving as much sunning: Exposing the plumage to the or group of species illustrates the general genetic diversity as possible. [10·76] sun, usually by spreading the wings and/ pattern of mortality. [8·5] subadult: A bird that has not yet reached or tail (often against the ground), fluffing sutures: Boundary lines along the junc- maturity; immature. [3·30] the feathers, and remaining motionless; tion between two bones; sutures are vis- subadult plumage: Any of the plumages sunning may aid Vitamin D production ible in the skull of a young bird. [4·10] worn by young birds before they reach and remove ectoparasites, as well as sweeping: A foraging technique in which their definitive plumages (those of a ma- warm the bird. [3·22] a bird pursues insects in flight, captur- ture bird). [3·30] superciliary line: See eyebrow stripe. ing them in midair in its mouth; used by subclavian arteries: Two arteries (left and [1·7, 2·10] swifts, nighthawks, and many swallows, right) that carry blood to the front limbs supplemental plumage: In the Hum- among others. [6·44] (wings). [4·82, 4·84] phrey-Parkes system of nomenclature, a syllable: In a bird song, a note or cluster subclavian veins: Paired veins (left and plumage that may occur in addition to the of notes that forms a unit that is repeated. right) that carry blood from the front limbs basic and alternate plumages; it is found [7·9] (wings); on each side, they merge with in just a few species, such as ptarmigans symbiosis: An ecological relationship the jugular and pectoral veins to form the and certain buntings. [3·33] between organisms of two different spe- cranial vena cava, which returns blood to supracoracoideus: The powerful flight cies that live in close association with one the heart. [4·83] muscle (also called the supracoracoid another; types of symbioses include mu- subelliptical: Describes an egg that is muscle) that raises a bird’s wing; it also tualism, commensalism, and parasitism longer than an elliptical egg, with a slight slows down the wing at the end of the (see separate entries for each). [9·132] bulge to the sides. [8·73] downstroke and accelerates it at the be- sympathetic ganglion: A clump of cell ginning of the upstroke. [5·6] suboscines: Members of Suborder bodies of sympathetic neurons that arise Tyranni, one of the two large suborders supracoracoid foramen: See foramen in the spinal cord. A chain of sympathetic of Order Passeriformes (perching birds). triosseum. [4·20, 5·6] ganglia runs the length of the spinal cord The suboscines have less complex voice surface diving: Diving under water from a near its ventral surface. [4·44] boxes than members of the other pas- swimming position on the water’s surface; sympathetic system: The part of the au- serine order, Passeri, and thus must sing performed by birds such as diving ducks, tonomic nervous system that functions simpler songs. Includes birds such as ty- grebes, cormorants, and loons. [6·45] under conditions of stress; the sym- rant flycatchers, antbirds, manakins, and surface-to-volume ratio: An individual’s pathetic system prepares the body for cotingas. [7·25] surface area (through which heat is lost) “fight or flight” by increasing the heart subsong: Practice singing (somewhat like divided by its volume (which generates rate, breathing rate, and blood pressure. human infant babbling) that begins short- heat); the larger the ratio, the faster an in- It consists of nerves that leave the spinal ly after songbirds leave the nest. Subsong dividual will lose heat. Smaller birds have cord from the thoracic and lumbar re- usually is quiet, garbled, and rambling larger ratios than larger birds, and thus to gions. [4·44] compared to adult song; it contains some compensate they must have higher meta- synapse: A small gap between nerve of the same elements, but they are strung bolic rates and must consume relatively cells, across which nerve impulses travel. together in odd ways. It gradually comes more food. Also called surface area-to- [4·32] to resemble adult song. [7·27] volume ratio. [3·11, 4·146] synchronous hatching: Pattern of hatch- subspecies: A subset of a species, usu- surrogate: In conservation biology, a ing in which all the eggs of a single clutch ally in a particular geographic area, that stand-in for another individual or spe- hatch at about the same time (on the same contains individuals that are morpho- cies. For example, researchers practiced day), resulting in a brood of young all the logically distinct from other individuals releasing techniques for the highly en- same size and age. This hatching pattern of the same species, but are still capable dangered California Condor by first using occurs when incubation is delayed until of interbreeding with those other indi- the biologically similar but less endan- the last egg is laid. Because development viduals. Also called a race. [1·55] gered Andean Condor as a surrogate. of laid eggs does not begin until they are sugarbirds: A family (Promeropidae, [10·102] warmed, all the embryos begin to de- 2 species) of long-billed, long-tailed, survival rate: 1. The proportion of indi- velop at the same time and are ready to nectar feeders that specialize on Pro- viduals in a population that survive for hatch at the same time. [6·88] tea plants on the mountainous slopes of a particular interval of time—usually a syndactyl feet: Foot arrangement in South Africa. [1·85] year. [9·64] 2. The chance that a par- which the hallux points backward and sunbirds: A family (Nectariniidae, 124 ticular individual will survive a given toes two, three, and four point forward, species) of tiny birds with long, slender, period of time, usually one year. For ex- with toes two and three (the inner and down-curved bills; they feed on nectar ample, an adult Royal Albatross has an middle toes) fused for much of their and insects and serve as pollinators of annual survival rate of 95 percent. [8·3] length. Found in many kingfishers and flowers. Sunbirds are similar to hum- survivorship: The proportion of individu- hornbills. [1·21] mingbirds, but do not hover while feed- als from a cohort (a group containing in- synsacrum: The segment of the vertebral ing. Many males are brilliantly colored. dividuals born during the same period of column of birds that is formed by the fu- Most species inhabit Africa, but some are time) that survive to a given age. [8·5] sion of some thoracic vertebrae with all of found in tropical parts of the other Old survivorship curve: A graph of the pro- the lumbar, all of the sacral, and the first World regions. [1·86] portions of individuals in a cohort (a few caudal vertebrae; the synsacrum is sun compass: A mechanism by which birds group containing individuals born during in turn fused on either side with the ilium and some other animals use the position the same period of time) that remain alive bones of the pelvis. [4·18, 4·24]

Handbook of Bird Biology 56 Syr – Tip Glossary syringeal muscles: The muscles of the temporal fovea: Area—in the posterior brae above and to the sternum (breast- syrinx; they allow the syrinx to change quadrant of the retina of hawks and oth- bone) below. It forms a flexible but strong shape and thus to produce different types er fast-flying diurnal avian predators— enclosure for the heart, liver, and lungs, of sounds. [4·93] where the cones are most concentrated as well as for the thoracic air sacs of birds. syrinx: The sound-producing organ of and the neural layer (the nerves from the [4·19] birds; located at the point where the tra- rods and cones, which overlie the rods thoracic ducts: Paired lymph ducts that chea divides to form the two bronchi. and cones and block some light) is the collect the products of fat digestion from [4·93, 7·38] thinnest, and thus vision is the sharpest. the intestinal lymph trunk (coming from The temporal foveae provide sharp bin- systemic circulation: Portion of the cir- the small intestine; see separate entry); ocular views of the area in front of the the thoracic ducts run along the surface culatory system that carries blood from bird. [4·49, 4·53] the heart to the body tissues (excluding of the aorta and eventually deliver their the lungs) and back to the heart. [4·77] tendon: A band of fibrous connective tis- contents to the venous system at the cra- sue that attaches muscles to bones, and/ nial vena cavae just before these large or binds many muscle fibers together to veins enter the heart. [4·89] form a skeletal muscle. [4·6] thoracic vertebrae: The vertebrae of T teratorns: Giant, soaring, scavenging the thorax (chest) region; they articulate taiga: See boreal forest. [9·116] birds of the New World (subfamily Tera- with the ribs and thus form part of the rib tail bone: See pygostyle. [4·18] tornithinae) whose fossils date from the cage. In birds, some thoracic vertebrae tapaculos: A Neotropical suboscine fam- late Tertiary period; teratorns became are fused with lumbar, sacral, and some ily (Rhino-cryptidae, 52 species) related extinct around 10,000 years ago when caudal vertebrae to form the synsacrum. to antbirds; tapaculos have cocked tails many large mammals were wiped out. [4·15, 4·18] and frequent more southerly, open, and The largest species had a wingspan ap- thrombocytes: Nucleated blood cells dry areas than do antbirds. [1·79] proaching 19.4 feet (5.9 m). [10·8] that resemble red blood cells, but are tarsals: Ankle bones. In humans, they termitary: An active termite nest. [8·19] more dense and complex and are highly remain distinct; in birds, the proximal territory: A defended area. [6·22] specialized to carry out blood clotting. They are not present in mammals, which tarsals fuse with the tibia to form the tib- tertiary consumers: Level in a food chain iotarsus, and the distal tarsals fuse with instead have platelets (not found in birds) or web that consists of organisms—usu- to carry out blood clotting. [4·88] the metatarsals (instep bones) to form the ally animals—that directly eat secondary tarsometatarsus. [1·14, 4·25] consumers. [9·123] thrust: In a bird, the portion of the force tarsometatarsus: The long bone sup- generated by the flapping wings that pro- testes (singular, testis): The male gonads; pels the bird forward (in the direction the porting the upper section of the bird foot; one testis lies at the cranial end of each the tarsometatarsus is formed by a fusion bird is moving, which is always the direc- kidney. The testes produce sperm as well tion opposite to drag). [5·18] of the distal tarsals (ankle bones) and the as the male sex hormone, testosterone, metatarsals (instep bones). Also called and in some species, a female sex hor- thyroid glands: Paired endocrine glands simply the metatarsus. [1·14] mone, estrogen. [4·127] at the base of the neck. Under the con- trol of the anterior pituitary, the thyroid tarsus: The upper section of the avian foot, thecodonts: A diverse group of reptiles between the heel and the toes. [1·15] glands secrete the hormones thyroxin from the early Mesozoic, all of which have and triiodothyronine, which regulate the taste buds: Simple structures, usually a diapsid skull, teeth set in sockets, and an annual increase in gonad size, sperm and embedded in the epithelium of the oral antorbital fenestra (an opening on each egg production, the growth and pigmen- cavity and the tongue, that are the recep- side of the skull in front of the eye socket). tation of feathers, and molting. [4·74] tors for taste sensations in all vertebrates. Also known as basal archosaurs or pseu- Humans have numerous taste buds locat- dosuchian thecodonts. [E·8, E·33] tibia: In humans, the larger of the two ed on the tongue, but birds have few taste lower leg bones; in birds, the tibia fuses thermal: A rising column or large bubble with several tarsal (ankle) bones to form buds, which are located primarily on the of warm air that results from differential roof of the mouth or deep in the oral cav- the tibiotarsus, which supports the crus heating of land surfaces by the sun; ther- (lower leg). [1·14] ity, with none on the tongue. [4·65] mals develop over areas that are darker taxonomy: The classification of organ- than surrounding areas, and over south- tibiotarsus: Bone supporting the lower leg isms—assigning names and relation- facing slopes. [5·39] (crus) of birds; the tibiotarsus is formed by ships. [1·32] the fusion of the tibia with the proximal thermal soaring: A type of static soaring tarsal (ankle) bones. [1·14, 4·25] tectorial membrane: The upper mem- (see separate entry) in which birds use the brane of the cochlear duct in the inner rising air in thermals (rising columns of tickbirds: See oxpeckers. [1·87] ear. Sound waves set the lower mem- warm air) to propel themselves upward, time compensation: In the course of brane—called the basilar papilla in circling higher and higher with little en- orientation or navigation, making allow- birds—into motion, moving its hair cells ergy expenditure of their own. Once high ances for the changes in the position of against the tectorial membrane and trig- in the air, they can glide out of the thermal the sun (or other celestial bodies) in the gering nerve impulses in the hair cells, and across the countryside in whatever sky throughout the day. [5·84] which are sent to the brain in the process direction they wish to travel. [5·39] tinamous: Primitive, grouse-like birds of sound perception. [4·59] thigh: The upper leg, supported by the (family Tinamidae, 46 species) whose ee- telencephalon: The anterior portion of femur. [1·14] rie calls haunt both forests and pampas of the forebrain; it contains the olfactory thoracic air sacs: Two pairs of air sacs South and Central America; tinamous are lobes. [4·36] in the chest region of birds, one pair (a the only ratites capable of flight. [7·24] temperate: Moderate; describes a cli- cranial and caudal sac) located on each tip vortex: A type of turbulence created mate free from extreme heat and cold, side of the body. [4·101] at the tip of a wing; it consists of air cur- but experiencing some of both; generally thoracic cage: The rib cage; it consists of rents spiraling off the wing tip behind a found in the middle latitudes. [1·69] the ribs connected to the thoracic verte- bird or other flying object. These eddies Cornell Laboratory of Ornithology Glossary Tis – Tym 57 sometimes may be seen as trails of white transverse plane: A vertical plane through each nostril is separate, but in others the behind the wing tips of large airplanes as an organism, dividing the body into cra- two adjacent tubes fuse to form a double they take off or land. [5·34] nial and caudal portions. [1·4] tube—within which the tubes coming tissue: An aggregation of cells with related trees-down theory (of the origin of avian from each nostril may remain separate and often very similar characteristics; ex- flight): See arboreal theory. [E·18] or may be partially fused. Tubular nares are found in the Procellariiformes, an amples include muscle tissue and bone. trial-and-error learning: Learning to as- [4·2] order of pelagic seabirds that excrete sociate one’s own behaviors with either large amounts of salty fluid through their tissue fluid: Blood plasma that has moved a reward or punishment; behaviors that nostrils. The tube is thought to reduce out of blood cells at the capillary beds, result in reward are repeated, and those heat and airflow at the nostril (where the and is in the minute spaces between the that result in punishment are abandoned. nasal cavity opens into the upper beak), body cells. Oxygen and nutrients diffuse [6·10] allowing the salty fluid to flow down the out of it to the body cells, and carbon di- trigeminal nerve: The fifth cranial nerve; beak and away from the nostril before oxide and other wastes diffuse in. Tissue it divides into the ophthalmic, maxillary, evaporating. Thus the salts remaining fluid eventually returns to the blood sys- and mandibular nerves (see separate en- after evaporation are less likely to clog tem, either by diffusing directly back into tries) after exiting the brain. [4·41] the nostril and salt gland than if evapora- a blood capillary near the venous side, or trill: A phrase or song that consists of a tion occurred right at the nostril. [3·39, by diffusing into a lymphatic capillary (in 3·40] which it is called lymph), which carries single note or cluster of notes (a syllable) it to larger lymph vessels and eventually that is repeated many times in rapid suc- tufa: A porous, white limestone formed into the venous system. [4·87] cession. [2·16] under water when springs or rainwater carry dissolved calcium into lakes that tongue bone: See entoglossal. [4·14] triosseal canal: See foramen triosseum. [4·20, 5·6] have a high concentration of bicarbonate torpor: A profound state of sleep in ions (found in baking soda). When the trochlear nerve: The fourth cranial nerve; which the body temperature drops and two solutions mix they form calcium car- it carries motor output from the brain to a consequently all metabolic processes bonate, which precipitates (comes out single eye muscle. [4·41] and stimulus-reaction processes slow of solution), creating tufa—which often down. Used by swifts, hummingbirds, trophic: Pertaining to feeding. [9·123] accumulates into underwater towers or nighthawks, and some other birds to trophic levels: The different levels of food crusts covering the lake bottom. Mono conserve energy when food resources are production or consumption within a food Lake, in California, is well known for its unavailable. Birds may enter torpor on a chain or web; for example, producers, tufa formations, which became exposed nightly basis, or for extended periods up primary consumers, secondary consum- when the lake’s water level dropped dra- to an entire season of cold weather. Also ers, and decomposers. [9·123] matically after water was diverted for use called hibernation. [4·154] trophic structure: Organizational in the Los Angeles area. [8·60] trachea: The windpipe; the tube con- scheme of a community that is based on tundra: See arctic tundra or alpine tun- ducting air from the larynx to the lungs. the feeding relationships between organ- dra. [9·114] [4·92] isms: species are assigned to different tro- turacos: African relatives of cuckoos, tu- tracheal bulla: An expanded sac on one phic levels according to what they eat racos are noisy and arboreal, and tend side of the lower end of the trachea in and what eats them (see trophic levels). to run squirrel-like along branches. The males of many duck species; it is thought [9·123] plumage of turacos is soft green (one to modify the sounds produced by the Tropical Marine region: The major faunal of the few green pigments known from syrinx. [4·92] region of the seas that includes the warm birds), blue, or gray, often with bright red track: A flying bird’s actual direction of equatorial waters between the Subtropi- (or in a few species, purple) on the wings. movement with respect to the ground; due cal Convergences of both hemispheres. The 18 species of turacos, together with to crosswinds, a bird’s track may not be the [1·99, 1·101] plantain-eaters and go-away-birds, make same as its heading. Also applies to aircraft true navigation: The ability to take the up the family Musophagidae. [1·86] and other flying organisms. [5·68] proper course toward a specific goal, turbulence: A disorderly flow of air; tur- tracts: Bundles of axons and their my- even from completely unfamiliar sites at bulence may interfere with smooth air- elin sheaths within the central nervous vast distances from known areas. [5·83] flow over a bird’s wings and thus may system; collectively, tracts form the white trumpeters: A family (Psophiidae, 3 spe- disrupt flight. [5·13] matter, found in the outer portions of the cies) of large, hump-backed, chicken- tutor song: A song played to a bird in the spinal cord and the inner areas of the like birds that inhabit the rain forests and laboratory during experiments on song brain. [4·36] floodplains of Amazonia. Trumpeters learning. [7·26] transequatorial migrants: Birds that mi- forage on the ground in flocks, but roost tympanic canal: The lower perilymph- grate back and forth across the equator to and nest colonially in trees. They eat both filled canal inside the bony cochlea of take advantage of spring and summer in plants and animals (including reptiles the inner ear; the tympanic canal is con- both hemispheres. [1·103] and amphibians, and even carrion), and nected to the (upper) vestibular canal at transitive inference: The ability to un- sometimes follow army ants for the ar- one end. Pressure waves in the perilymph derstand and infer linear relationships thropod prey they disturb. [1·74] of the two canals are transmitted to the among individuals. For example, the tubenoses: Birds in the order Procellari- endolymph of the cochlear duct between ability to understand that “if A is dom- iformes, including albatrosses, shearwa- them, setting the (in birds) basilar papilla inant to B, and B is dominant to me, then ters, fulmars, petrels, and storm-petrels, in motion and stimulating the sensory A is dominant to me.” [6·18] all of which have tubular nares (see sepa- hair cells. [4·57] translocation: Establishing individuals of rate entry). [3·40] tympanic membrane: A membrane that a species in an area in which that species tubular nares: Nares (nostrils) that open stretches tightly across the external ear formerly lived by importing individuals into a horny, tubular structure that sits on canal; the tympanic membrane vibrates from a different area. [10·89] top of the bill; in some species the tube for when struck by the pressure waves of a

Handbook of Bird Biology 58 Tym – Ven Glossary sound, and transmits those vibrations to each rib. Because each uncinate process tive birds such as Archaeopteryx. [E·4] the (in birds) columella. Also called the overlaps with the rib behind it, the pro- utriculus: One of the two membranous eardrum. [4·56] cesses help to strengthen the rib cage. chambers inside the bony vestibule of the tympaniform membranes: Flexible Found only in birds. [4·18] inner ear (the other is the sacculus); these membranes that stretch between suc- understory vegetation: The shrubs and chambers contain hair cells and dense cessive cartilaginous rings of the syrinx, shorter trees in a forest; the understory crystals called statoconia, both of which allowing the syrinx to change shape (in vegetation grows below the level of the are embedded in a gelatinous material response to contractions of the syringeal main canopy and mid-story vegeta- that is surrounded by endolymph. The muscles) and thus to produce different tion (see separate entry), and above the statoconia and hair cells perceive the po- types of sounds. [4·93] ground vegetation. [9·93] sition of the head with respect to gravity Tyranni: One of the two large suborders of undertail coverts: The short contour feath- (see entry for statoconia). [4·58] Order Passeriformes (perching birds). The ers that cover the bases of the rectrices on Tyranni (also called suboscines) have less the ventral side of the tail. [1·6, 1·12] complex voice boxes and thus sing sim- underwing coverts: The feathers lining pler songs than members of the other pas- V the underside of the wing, from the lead- vagina: In birds, the short, lower section serine order, Passeri (also called oscines). ing edge of the wing to the flight feathers. of the oviduct, after the shell gland; the Suborder Tyranni includes birds such as [1·12, 1·13] vagina opens into the left side of the clo- tyrant flycatchers, antbirds, woodcreep- aca. [4·130] ers, ovenbirds, tapaculos, manakins, and upper beak: The upper half of the beak; cotingas. [7·25] also called the upper jaw. [1·6] vagus nerve: The tenth cranial nerve; it carries sensory and motor information Tyrannidae: Extremely diverse New upper jaw: See upper beak. [1·6] between the brain and the pharynx, lar- World suboscine family containing uppertail coverts: The short contour feath- ynx, heart, lungs, gizzard, liver, and in- nearly 400 species; members of the fam- ers that cover the bases of the rectrices on testines. [4·42] ily Tyrannidae are commonly known as the dorsal side of the tail. [1·12, 1·13] tyrant flycatchers. No other bird family upwelling: A process by which cold, nu- vane: The broad, flat surface on each side has more species. [1·80] trient-rich water from the lower ocean of the shaft of a contour feather; the vanes are formed by the interlocking of adja- tyrant flycatchers: The nearly 400 spe- depths rises to the surface. Upwelling cent barbs. [3·3] cies of the diverse New World suboscine may occur where certain ocean currents family Tyrannidae. [1·80] meet, or where winds blowing away from vangas: A diverse songbird family (Vangi- the coast move the surface water away dae, 14 species) of shrike-like birds. Most from shore, causing water from below to vanga species are gregarious and noisy, rise to replace it. [1·103, 1·105] gleaning insects and other small animals U ureters: Two tubes, one leading from as they move through the trees in groups. ulna: The larger and thicker of the two each kidney, that carry urine from the [1·88] bones that make up the antebrachium. kidneys to the (in birds) cloaca. [4·125] vasa deferentia: See deferent ducts. [1·9] uric acid: A complex but less toxic mol- [4·128] ulnare: The larger of the two wrist bones ecule to which the liver of birds converts vector: 1. A quantity with both magnitude of birds; the ulnare consists of several the highly toxic nitrogenous waste prod- (size) and direction; for example, a bird’s fused carpals. [1·9] ucts of protein metabolism. Compared flight at 25 miles per hour to the south ultimate: In biology, pertaining to long- to urea, the nitrogenous waste product is considered a vector. A vector is often term causes; evolutionary. Ultimate of mammals, uric acid requires more en- shown as an arrow whose length repre- questions about a behavior, for ex- ergy to produce, but it is less soluble in sents magnitude and whose orientation ample, look into the factors that might water and more highly concentrated, al- indicates compass direction. [5·68] 2. have caused the particular behavior to lowing birds to excrete more toxic waste An organism that acts as a carrier, trans- evolve—why it evolved. For comparison, per molecule excreted, while minimizing mitting pathogens (viruses, bacteria, and see proximate. [6·3] water loss. [4·125] so on) from one individual to another. For ultimate factors: In biology, the variables urinary bladder: Sac that stores urine be- example, certain mosquitoes are vectors affecting the survival and reproduction fore it is excreted; among birds, the uri- because they carry malaria from one per- of a population and its ancestors, and nary bladder is pre-sent only in the South son to another. [9·70] therefore influencing the evolution of American rheas. [4·125] vector navigation: Finding one’s way by certain traits, behaviors, or physiological urinary system: See excretory system. adding vectors composed of compass processes, or the timing of such things. [4·125] direction and distance in order to deter- For example, the food supply at different mine the straight-line path to a desired times of the year is a major ultimate fac- urogenital system: The combination of destination. [5·79] tor that affects the timing of breeding in two organ systems: the urinary (excreto- ry) system, which removes toxic nitrog- vein: A vessel that carries blood toward birds. For comparison, see proximate the heart. All veins except the pulmonary factors. [8·11] enous wastes from the blood, and the genital (reproductive) system. The two vein and its tributaries carry oxygen-poor ultimobranchial glands: Two small, light- systems often are considered together blood. [4·81, 4·84] colored glands located near the parathy- because they are located close together vena cava: See caval veins or cranial vena roid glands, usually on an artery; the in the body, and because they may share cavae or caudal vena cava. [4·84] ultimobranchial glands secrete the pro- some structures (such as the cloaca, in venous ring: A ring of veins formed by the tein calcitonin, which lowers the blood birds). [4·124] calcium concentration. [4·74] left and right renal portal veins of birds; the uropygial gland: See oil gland. [3·20] venous ring connects the various lobes of uncinate process: A flattened, hook- urvogel: General term (used primarily by the two kidneys, and is part of the avian shaped extension of bone that projects renal portal system. [4·83] caudally from the vertebral segment of the Germans) that refers to early, primi-

Cornell Laboratory of Ornithology Glossary Ven – Wet 59 vent: The opening of the cloaca to the containing two membranous chambers, separate entry), which gives rigidity to exterior of the body. In birds, the vent is the utriculus and sacculus, which contain the eyeball. [4·48] the only posterior opening, and thus it hair cells that perceive the position of the vocal repertoire: All the different types releases feces from the digestive system, head with respect to gravity. The vestibule of vocalizations that are produced by uric acid from the excretory system, and is part of the bony labyrinth of the inner an individual bird, including both songs sperm or eggs from the reproductive sys- ear and thus contains perilymph. [4·48] and calls; for comparison, see song rep- tem. [4·123] vestibulocochlear nerve: The eighth cra- ertoire. [7·10] ventral: Toward the belly of an organism. nial nerve; the vestibulocochlear nerve vocal signals: Vocalizations that have [1·4] carries sensory input for balance and evolved for a specific function. [7·11] ventricles: The two thick-walled, muscu- hearing. Its vestibular (balance) portion carries sensory input from the semicircu- voluntary muscles: See skeletal muscles. lar, posterior chambers of the heart; the [4·26] ventricles receive blood from the atria lar canals and vestibule of the inner ear, and then pump it to other parts of the and its cochlear (hearing) portion carries body: the right ventricle pumps blood sensations from the hair cells of the inner to the lungs and the left ventricle pumps ear. Formerly called the auditory, acous- W blood to the body arterial system. [4·77] tic, or statoacoustic nerve. [4·42] warm-blooded: See endothermic. [1·1] venules: Small veins that carry blood vestigial structure: A structure that has warm front: The interface between a from the capillaries to larger veins. [4·81, little or no apparent function and is warm air mass and the cold air mass it 4·84] thought to be present in an organism is overtaking; the warm air tends to push only because it has been inherited from vertebrae (singular, vertebra): The indi- up over the denser cold air and become an ancestor. Many vestigial structures are cooler, which results in the formation vidual bones that form the vertebral col- reduced in size from the ancestral (and umn (backbone). [4·14] of clouds and eventually precipitation. functional) condition, and presumably [5·69] vertebral canal: A long tube that runs the are in the process of being eliminated length of the back, inside the backbone; through natural selection. A vestigial water birds: See aquatic birds. [1·65] the vertebral canal is formed by the ver- structure in humans is the remains of a waterfowl: Ducks, geese, and swans; tebral arches of successive vertebrae. The nictitating membrane in the inner corner family Anatidae. [1·65] vertebral canal protects the spinal cord, of each eye—in reptiles, birds, and some wattlebirds: Common name for mem- which runs inside it from the brain to the other mammals the nictitating membrane bers of the family Callaeidae, which tail. [4·38] is functional. [3·46] consists of two living species (the Sad- vertebral column: Commonly called the viduines: Refers to members of the Af- dleback and Kokako) and one extinct backbone or spine, the vertebral column rican genus Vidua, which consists of species (the Huia). Wattlebirds are for- is a series of complex, uniquely articulat- whydahs and indigobirds. Viduines are est-dwelling songbirds endemic to New ing or rigidly fused vertebrae that provide colorful seed-eaters that are brood para- Zealand; they are named for the pair of support for the head, neck, ribs, back, sites on other members of their family, Es- colorful, fleshy wattles at the corners of and tail. [4·14] trildidae; they are known for the intricate their mouths. [1·97] vertebral rib: The dorsal portion of each patterns inside the mouths of their nest- wattles: Ornamental flaps or folds of thoracic rib; the vertebral rib articulates lings, which strikingly resemble those of skin that dangle from the head or neck of with the thoracic vertebra dorsally and their host’s nestlings. [8·142] some birds. Wattles are found in turkeys, with the sternal rib ventrally. [4·18] villi (singular, villus): Minute fingerlike pheasants, cotingas, wattlebirds, and vertebrates: Animals that have a back- projections. Villi often are produced by many others. [3·48] bone: birds, fish, amphibians, reptiles, folds in a tissue lining an organ—as in waxbills: Together with whydahs, indi- and mammals. [1·1] the inner surface of the small intestine of gobirds, and numerous related species, vertebrates, where the presence of villi vertical migration: Type of migration in these colorful seed-eaters of the Old greatly increases the surface area avail- which animals move seasonally up and World form the large family Estrildidae able for absorbing nutrients. [4·121] down mountainsides. [5·53] (159 species). The Zebra Finch, a well- visual field: See field of view. [4·51] known cage bird, is a waxbill. [1·83] vestibular canal: The upper perilymph- filled canal inside the bony cochlea vitelline diverticulum: Small pouch of weavers: A diverse Old World fam- of the inner ear; the vestibular canal is tissue at the junction of the jejunum and ily (Ploceidae, 117 species) of colorful connected to the lower perilymph-filled ileum of the small intestine in many birds; seed-eaters; weavers are found primar- canal, the tympanic canal, at one end. the vitelline diverticulum is a remnant of ily in open tropical areas and are named Pressure waves in the perilymph of the the yolk sac of the embryo. [4·122] for the ability of many species to weave two canals are transmitted to the en- vitelline membrane: In avian biology, large, complex nests. [1·83, 8·53] dolymph of the cochlear duct between the transparent membrane surrounding well: See sap well. [9·128] them, setting the (in birds) basilar papilla and holding together the yolk of an egg. western montane forest: North Amer- in motion and stimulating the sensory [8·62] ican coniferous forest ecosystem that is hair cells. [4·57] vitreous body: Clear, gelatinous material dominated by spruce and fir trees and vestibular window: A soft, oval region on filling the vitreous chamber (see sepa- is found along the North Pacific slope the bony cochlea, to which (in birds) the rate entry) inside the eyeball. The vitre- and in the Cascades, Sierra Nevada, and columella of the middle ear is attached. ous body gives rigidity to the eyeball and Rocky Mountains; western montane for- Movement of the columella moves the helps to maintain its shape. [4·48] est hosts few breeding birds. [9·116] vestibular window, sending pressure vitreous chamber: The largest chamber wetland mitigation: Compensation that waves through the fluid (perilymph) in- inside the eyeball; the vitreous chamber the recipient of a permit to fill or otherwise side the cochlea. Formerly called the oval is located on the posterior side of the eye- destroy wetland areas is legally required window. [4·57] ball and is filled with a clear, gelatinous to provide, to make up for the wetland vestibule: Bony structure of the inner ear material called the vitreous body (see area lost. Compensation may consist of Handbook of Bird Biology 60 Whi –Zyg Glossary restoring or re-creating other wetland ar- ines). Viduines are colorful seed-eaters adult birds helping the parents to tend eas of equal or greater size compared to that are brood parasites on other mem- the nest. [1·85] the wetland area destroyed, or may be a bers of their family, Estrildidae; they are woodswallows: A family (Artamidae, cash payment into a mitigation bank—a known for the intricate patterns inside 14 species) of small, chunky songbirds fund managed by a public agency or the mouths of their nestlings, which with a graceful, swallow-like flight; private conservation group that carries strikingly resemble those of their host’s woodswallows inhabit the Australasian out large-scale wetland restoration proj- nestlings. [8·142] and Oriental regions and are known for ects using money accumulated from a wick: In avian biology, a beakful of veg- huddling together on branches or in tree large number of small-impact projects. etation that is dipped in mud and then cavities in groups of up to 50 or more [10·93] used to transport the mud to a nest that individuals. [1·96] whisker stripe: A distinctively colored is under construction; usually the entire wood-warblers: A family (Parulidae, 115 stripe in the malar region of birds; also wick is added to the nest. [8·54] species) of small, insectivorous birds, called a mustache stripe or malar stripe. wing-flashing: A foraging technique many of which are colorful, found in the [1·8, 2·10] used by some herons, egrets, and storks New World. Many wood-warblers are whistle: A song or phrase in which clear in which a bird wading through the water Neotropical migrants. Also called New tones are produced one pitch at a time. quickly raises or brings forward one or World warblers. [1·71] [2·16] both wings, apparently to frighten prey whistlers: Large, round-headed song- out of hiding or to provide a shady place birds with slightly hooked bills; whistlers where unsuspecting prey may try to hide, are known for their explosive, often- thus bringing the prey within the bird’s Y yolk: In avian biology, the familiar, yel- beautiful, songs. The nearly 40 species reach. [6·46] low portion of an egg; the yolk contains of whistlers, along with shrike-thrushes, wing loading: The ratio of body weight to nearly all of the lipids (fat) and most of pitohuis, and a few related species, form wing area; wing loading is a measure of the protein needed by the developing the family Pachycephalidae, which is how much “load” each unit area of wing embryo. The yolk is surrounded and found primarily in the Australasian re- must carry. [5·31] held together by the transparent vitelline gion. [1·95] wing pouch: A pocket under each wing membrane. [8·62] white blood cells: Unpigmented blood of male Sungrebes (Neotropical inhabit- yolking up: The deposition of yolk with- cells of several different types, all of ants of wooded streams); the wing pouch in the vitelline membrane in alternating which contain a nucleus and can leave is formed from a pleat of skin and is used bands of darker and lighter yolk; yolking the capillaries and move about in the to carry the young while the male flies or up is part of the egg-formation process spaces between the body cells. Many swims. [8·137] in birds and occurs before egg laying. types of white blood cells are important wing spurs: Bony outgrowths of the car- [8·63] in fighting infections. Also called leuko- pometacarpus (the main bone of the yolk sac: In avian biology, the extra-em- cytes. [4·88] avian manus); wing spurs are found in bryonic membrane inside the egg that white fibers: Muscle fibers that appear various birds such as cassowaries, plo- surrounds the yolk of the developing lighter in color than red fibers because vers, sheathbills, screamers, and jacanas. embryo. [8·65] they have fewer capillaries (and thus few- [4·22] er red blood cells) and less of the reddish winnowing: Part of a twilight territorial oxygen-transport molecule myoglobin; and courtship display given by both male white fibers also are larger in diameter and female Common Snipe. The birds cir- Z than red fibers. Muscles with many white cle high in the air and produce a series zooplankton: Microscopic animals and fibers are used for quick bursts of action, of rapid, pulsating, whistle-hums (called protozoa that float freely in aquatic en- but they cannot carry out sustained activ- winnows) by spreading the tail and div- vironments; zoo-plankton make up the ity because they produce energy anaero- ing at high speeds—causing the stiff outer “animal” portion of plankton. [1·103] bically (without oxygen)—so lactic acid tail feathers to vibrate. European Snipe Zugunruhe: See migratory restlessness. builds up quickly and causes fatigue. also winnow. [7·17] Muscles with many white fibers often are [5·60] called “white meat” (as in the breast of a winter plumage: See basic plumage. zygodactyl feet: Foot arrangement in turkey). [5·7] [3·33] which toes two and three point forward, white matter: Lighter-colored tissue wishbone: See furcula. [4·19] and the hallux and toe four point back- (compared to gray matter) that makes up woodcreepers: A Neotropical family ward. Found in woodpeckers, cuckoos, much of the brain and spinal cord. White (Dendrocolaptidae, 51 species) of simi- toucans, owls, Osprey, turacos, most par- matter consists of numerous tracts, which lar-looking, rust-colored, suboscines rots, and some other birds. [1·21] are bundles of nerve cell axons and their with a wide array of beak shapes; wood- zygote: The fertilized egg; the single- myelin sheaths, and is found in the outer creepers typically forage in tree bark celled product resulting from the union portions of the spinal cord and the inner much like the unrelated Brown Creeper of (1) the nucleus of the sperm cell from areas of the brain. [4·39] of North America. [1·78] the male and (2) the nucleus of the ovum white meat: See white fibers. [5·7] woodhoopoes: A family (Phoeniculi- (egg cell) from the female. In birds, fertili- dae, 8 species) of sociable African birds zation occurs in the infundibulum of the whydahs: Members, along with indi- oviduct. [4·133 gobirds, of the African genus Vidua with glossy, dark plumage and long tails; (members of this genus are called vidu- woodhoopoes nest in tree cavities and breed cooperatively—with additional

Cornell Laboratory of Ornithology About the Authors To ensure scientific accuracy and a high level of ex- Although plants rather than birds became his main interest, his appreciation of nature derives partly from memorable expe- pertise, one or more professional ornithologists with riences with birds: “Once when I was a teenager, on a stormy specific knowledge and research experience have November day, as I walked along a hedgerow, I was astonished authored each chapter of this book. To achieve maxi- to come across an isolated little tree crowded with silent birds I’d never seen before—Cedar Waxwings. Too exhausted to fly, mum clarity and readability, the twelve chapters were they merely shuffled down their perches a bit when I came carefully edited by Lab of Ornithology Education staff, close. And in March of some years, the air was alive day and resulting in better consistency and flow among the night with the sound of immense flocks of Canada Geese flying between the lake and the muddy cornfields around our house.” primary subjects. However, because each author has a Experiences such as these, he notes, quicken one’s appreciation unique writing style, students may notice some differ- of the vivacity and fascination of the natural world, and help us ences in tone and presentation among the chapters. to understand what conservationists are fighting for.

Birds and Humans: A Historical Perspective Introduction: The World of Birds Dr. Kevin J. McGowan Sandra G. Podulka, Marie Z. Eckhardt, and Daniel R. Otis Kevin J. McGowan is a research associate at the Cornell Lab of Sandra G. Podulka received a B.S. in Wildlife Biology from Ornithology. He received his B.S. and M.S. degrees in Zool- Cornell University and an M.S. in Zoology (Animal Behavior) ogy from Ohio State University, and a Ph.D. in Biology from from the University of Maryland, where she studied the function the University of South Florida, where he studied the social of song repertoires in Song Sparrows. After graduate school, development of Florida Scrub-Jays. In 1988, after working as a Sandy was a Research Technician at Cornell for Dr. Stephen non-game biologist for the Florida Game and Freshwater Fish T. Emlen, analyzing the social behavior of White-fronted Bee- Commission, he came to Cornell University, taking a position eaters, but discovered she wanted to spend more time sharing as Curator of the Ornithology and Mammalogy Collections in her love of nature with others, so turned toward environmental the Department of Ecology & Evolutionary Biology. In addition education. She worked at the Cayuga Nature Center for several to caring for the collections, he conducted research on crows years, and as an Adjunct Professor of Biology at Tompkins Cort- and taught classes in specimen preparation, field collecting, the land Community College from 1988 to 1996, teaching courses relationships among birds, and Neotropical canopy biology. He in biology and conservation. Since 1986, she has worked at moved to his present position at the Lab in 2001. the Cornell Lab of Ornithology, in research, writing, public Kevin’s primary research focuses on the behavioral ecology education, and editing—most recently as one of the editors of of birds. Currently he is studying the reproductive and social be- the Handbook of Bird Biology and the Home Study Course in havior of two crow species in central New York State and inves- Bird Biology. tigating the impact of West Nile virus on crow populations. He Sandy spent her childhood knee-deep in muddy ponds try- is an Elected Member of the American Ornithologists’ Union, ing to catch tadpoles and frogs, and roaming fields collecting Webmaster and a Director of the Federation of New York State butterflies—her first real love. She has enjoyed birds as long as Bird Clubs, and a member of the New York State Avian Records she can remember, but they did not take center stage until she Committee. Formerly, he was Secretary of the Ornithological took a summer ethology course at Cornell University from Dr. Societies of North America (OSNA), and editor of the Ornitho- Bill Dilger. He hauled his students out before dawn every morn- logical Newsletter, a bi-monthly OSNA publication. ing and taught them to recognize birds by their songs, and Sandy An avocational birder since childhood, Kevin has traveled has been listening to them ever since. She has participated in throughout North America and to Europe, Africa, and Central Christmas Bird Counts and Breeding Bird Atlases, and has trav- and South America to watch and study birds. eled to Costa Rica, Belize, Trinidad, and Peru to watch birds, but her favorite bird-watching site is her yard—which overlooks a beaver pond with an ever-changing cast of avian actors. A Guide to Bird Watching Marie Z. Eckhardt was a biologist in the Education Program at the Cornell Lab of Ornithology. Her academic background in Dr. Stephen W. Kress vertebrate zoology and experience as a museum consultant and Stephen W. Kress is vice-president for bird conservation for a collections manager at the New York State Museum provided the National Audubon Society and Manager of the Society’s a strong foundation for her varied contributions at the Lab. Maine Coast Seabird Sanctuaries. He also is ornithology Pro- Marie has been interested in animals and the natural world for gram Director for the Audubon Camp in Maine, an adjunct as long as she can remember—an interest she credits to early faculty member in the Wildlife Department at the University and regular museum visits. of Maine, Orono, and a Visiting Fellow at the Cornell Lab of Daniel R. Otis grew up on a farm in Upstate New York. Since Ornithology, where each year he teaches the popular course 1988 he has worked as a freelance editor, proofreader, and writ- Spring Field Ornithology. er for Living Bird and other Lab of Ornithology projects. Cur- As Director of Audubon’s Seabird Restoration Program, rently a Ph.D. student in Horticulture at Cornell, his research Steve advises and manages the development of techniques for focuses on conservation of the world's maple species and on re-establishing in Maine colonies of various seabirds, such as assessing the extent and effects of Norway maple invasiveness Atlantic Puffins; Leach’s Storm-Petrels; and Arctic, Common, on northeastern forests. and Roseate terns. In the Pacific region, he has studied the role 62 About the Authors of vocalizations in attracting endangered Dark-rumped Petrels J. B. Heiser is a Senior Lecturer in the Department of Ecology to artificial burrows in the Galápagos Islands, and Short-tailed and Evolutionary Biology at Cornell University. A vertebrate Albatross to decoys on Midway Island. He is author of many evolutionary ecologist, J. B. received his B.S. from Purdue books, including The National Audubon Society’s Birder’s University and his Ph. D. from Cornell University. For the past Handbook, The Bird Garden, and Project Puffin, as well as 35 years, he has taught a variety of courses in vertebrate com- the Golden Guide to Bird Life. He also has written numerous parative anatomy and ecology. scientific papers on seabird biology and conservation. J. B. began teaching on the Ithaca campus, but soon was During most of the year Steve lives on 33 acres of woods teaching field courses at the Shoals Marine Laboratory, an iso- and meadows near Ithaca, New York, where he manages his lated island facility in the Gulf of Maine that is cooperatively land for songbirds and works on methods for restoring popula- run by Cornell University and the University of New Hamp- tions of Northern Bobwhite. He spends summers on the Maine shire. Eventually he became Director of the Shoals program, a coast, continuing his lifelong interest in restoring nesting sea- position that he held for 15 years. He won the Clark Award for bird colonies. 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 Form and Function: The External Bird research focuses on the evolution and interrelationships of coral reef fish, he has considerable natural history experience Dr. George A. Clark, Jr. in tropical forests worldwide. In his travels he has watched birds George A. Clark, Jr. is Professor Emeritus of Ecology and Evo- (and fish) in every major biome and biogeographic region that lutionary Biology at the University of Connecticut in Storrs. the planet has to offer, but he still gets a thrill out of backyard He received his Ph.D. in Biology from Yale University, where birding in Upstate New York. 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. Birds on the Move: Flight and Migration His 200 publications reflect his research interests in the structure, behavior, distribution, and evolution of birds. He is Dr. Kenneth P. Able past president of the Association of Field Ornithologists, served Kenneth P. Able is Professor Emeritus of Biology in the Depart- as co-editor of the book Perspectives in Ornithology/ Essays ment of Biological Sciences at the State University of New York Presented for the Centennial of the American Ornithologists’ in Albany, where he had been a member of the faculty since Union, and has led educational field trips for groups to observe 1971. He received his B.S. and M.S. degrees from the University birds in North and South America, Europe, and Africa. of Louisville, and his Ph.D. from the University of Georgia. Ken’s A birder since his high school days in Pennsylvania, George research focuses on bird migration, particularly the mecha- now resides in Vermont and enjoys seeking birds by walking or nisms of orientation and navigation. snowshoeing in the northern New England hills. Ken has been passionately interested in birds since child- hood. He has birded extensively across North America and in Mexico, Costa Rica, Peru, Spain, France, Germany, Italy, What’s Inside: Anatomy and Physiology Switzerland, the United Kingdom, Australia, New Zealand, and South Africa. His favorite destination is Australia, because the Dr. Howard E. Evans and Dr. J. B. Heiser avifauna is unique and the species fantastic. Howard E. Evans received both his B.S. and Ph.D. in Com- parative Anatomy from Cornell University, where he became a faculty member in the Veterinary College in 1950. There, he Evolution of Birds and Avian Flight taught gross anatomy of the horse and cow for seven years, and anatomy of the dog, bird, and fish for 36 years. He was Secretary Dr. Alan Feduccia of the college for 12 years and served as Chairman of Anatomy Alan Feduccia is S. K. Heninger Professor of Biology at the Uni- from 1976 to 1986, when he retired. He continues to teach a versity of North Carolina at Chapel Hill, where he has been for course on the literature and materials of natural history, and to 30 years. He received his Ph.D. at the University of Michigan, lecture in several other courses. where his thesis focused on the evolution of woodhewers and Howie’s research concerned the anatomy of reptiles and ovenbirds. birds, the replacement of teeth in fishes, the plant-induced cy- Alan’s career has focused on vertebrate evolution, the evo- clopis in sheep, fetal development of the dog, and anatomy lution of birds, and the tempo and mode of the evolution of of tropical fishes. His most recent works include the third edi- modern groups of birds. His interest in the origin of birds blos- tion of Miller’s Anatomy of the Dog (1993), the fifth edition of somed in the late 1970s when he wrote The Age of Birds for Guide to the Dissection of the Dog (2000, with Dr. Alexander Harvard University Press. In 1973 he wrote a rebuttal to the deLahunta), and the third edition of Anatomy of the Budgerigar theory of hot-blooded dinosaurs in Evolution, and since that and Other Birds (1996). He is co-editor of the Handbook of time has been involved in the debate on bird origins. His latest Avian Anatomy, published by the Nuttall Ornithological Club book, The Origin and Evolution of Birds (Yale University Press, of Harvard University. He has served as President of both the 1999), addresses many of the main issues in the bird evolution American and the World Association of Veterinary Anatomists, controversy, and takes what he calls the “ornithological” po- and has been an associate editor of the American Journal of sition—that birds were already arboreal when they evolved Anatomy and the Journal of Morphology. to fly, and that they evolved from a common ancestor with A native of New York City, Howie and his wife Erica have led dinosaurs, but not directly from them. natural history trips for Cornell Adult University to the Virgin Alan’s interest in birds began as a teenager. Later, as an Islands; Hawaii; Sapelo Island, Georgia; East and South Africa; undergraduate student at Louisiana State University, he had the Papua, New Guinea; and Antarctica. He has lectured in China, good fortune to participate in “bird” expeditions to Honduras, Russia, Taiwan, Hungary, Switzerland, Germany, England, El Salvador, and Peru. He has maintained an interest in Neo- Thailand, Brazil, Mexico, and Japan. tropical birds ever since.

Cornell Laboratory of Ornithology About the Authors 63 Malte Andersson); Oxford University (with John Krebs); and Understanding Bird Behavior Cornell University (with Paul Sherman). David joined the Cor- Dr. John Alcock nell faculty in 1988 upon the retirement of Tom Cade, and his research on swallows worldwide, and Tree Swallows in Ithaca John Alcock is Regents’ Professor of Biology at Arizona State in particular, has thrived ever since. 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 Individuals, Populations, and Communi- Desert. Earlier research has focused on birds, however, and his textbook, Animal Behavior, An Evolutionary Approach, uses ties: The Ecology of Birds many bird examples to illustrate all aspects of the modern study of behavior. Dr. Stanley A. Temple John began bird watching at age five (bird number one was Stanley A. Temple is the Beers-Bascom Professor of Conserva- the Mallard), and seeing a good bird still boosts his heart rate. In tion in the Department of Wildlife Ecology at the University attempts to keep his heart rate up, he has visited Costa Rica, Ec- of Wisconsin in Madison. He is also the Chair of the graduate uador, Argentina, Australia, and several countries in Europe. program in Conservation Biology and Sustainable Develop- ment in the Institute for Environmental Studies at Madison. A quintessential Cornellian, Stan earned his B.S., M.S., and Ph.D. Vocal Behavior all at Cornell, and for years has served as a member of the Lab of Ornithology’s Administrative Board. Dr. Donald E. Kroodsma Stan’s professional activities focus on avian ecology and bird Birds enter our lives in different ways. Some of us have been conservation, with a special emphasis on endangered species. bird-crazy as long as we can remember, but others discovered He and his students have worked with some of the world’s birds later. I was a late-comer, as birds grabbed me from the most endangered birds, including Peregrine Falcons, California chemistry lab during my last year of college. How grateful I am Condors, Whooping Cranes, and dozens of endangered species that they have never let go. Immediately after college I took two endemic to islands around the world. To date, none of those summer bird courses from the famed Olin Sewall Pettingill, species has become extinct, and most are doing significantly who was then director of the Cornell Lab of Ornithology but better as a result of his work. taught at the University of Michigan field station in Pellston, Stan has been interested in birds as long as he can remem- Michigan. He put a tape recorder, headphones, and parabolic ber. Birds of prey have always been among his favorites, and microphone in my hands and told me to “go out and tape record he has been a falconer for 45 years. He feels fortunate to have some birds.” In doing so, he changed my life. I went to graduate incorporated all of his ornithological pleasures into his profes- school at Oregon State to study how young wrens learn their sional life. 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 Bird Conservation University of Massachusetts, Amherst. My time since college has been spent reveling in the who, what, when, where, how, Dr. John W. Fitzpatrick and why of bird song. 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 Nests, Eggs, and Young: Biological Station, a private ecological research foundation in central Florida, from 1988 through August 1995, and was Breeding Biology of Birds Curator of Birds at the Field Museum of Natural History (Chi- Dr. David W. Winkler cago) from 1978 to 1989. He is a Fellow of the American Or- nithologists’ Union (AOU). In 1985 the AOU awarded him its David W. Winkler was born and raised in Sacramento, Cali- highest research honor, the Brewster Award, for his book and fornia. Unlike the rest of his family, he was a naturalist from numerous research articles (co-authored with Glen Woolfen- about age four. After a progression of enthusiastic interest in den) on demography, social behavior, and conservation of the butterflies, wildflowers, and herps, he finally settled on birds endangered Florida Scrub-Jay. Fitz has led many expeditions in his early teens. He learned and studied local birds alone for to remote areas of South America, especially the western Am- two years, when, at his first Christmas Bird Count, he ran into azonian basin and the Andean foothills. He has published nu- Rich Stallcup. David then spent the last two years of high school merous papers on Neotropical birds, including descriptions learning a great deal from Rich about the birds of California. of seven bird species new to science. Co-author of the book While in high school, an American Birds article introduced Neotropical Birds: Ecology and Conservation (University of David to The Herring Gull’s World by Niko Tinbergen, and he Chicago Press, 1996), he has been engaged in applying science carried out a senior English project on gull taxonomy within to real-world conservation issues throughout his career. Most a couple of years. By his freshman year at U. C. Davis, David recently, he helped design and implement a major network of was dreaming about all the species he might someday study, ecological preserves in central Florida by convening panels of as opposed to just see, for his list. scientific experts and by engaging county, state, and federal While attending U. C. Davis, David and friends secured agencies; nongovernmental organizations; and private industry National Science Foundation funding for an ecological study in the process. He serves on the national governing boards of of Mono Lake. This cemented his attachment to the Mono Ba- The Nature Conservancy, the National Audubon Society, and sin and its birdlife, and his friendship with David Gaines, with the Center for Biodiversity and Conservation at the American whom he co-founded the Mono Lake Committee in 1978. His Museum of Natural History. He is on two Endangered Species dissertation on the clutch sizes of California Gulls (with Frank Recovery Teams, including that of the world’s rarest bird, the Pitelka at Berkeley) at Mono and Great Salt Lakes was followed Hawaiian Crow. He enjoys watercolor painting, and has been by post-docs at the University of Gothenburg, Sweden (with a bird watcher since kindergarten.

Handbook of Bird Biology 64 About the Authors Illustrations 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 field biologist for both the Per- egrine Falcon and California Condor recovery programs. John’s work as a field biologist led to more serious devotion to illustrating birds. He has illustrated several books, includ- ing the National Geographic Society’s Third Edition of Birds of North America; Clark’s A Field Guide to the Raptors of Europe, the Middle East, and North Africa; and Skutch’s Birds Asleep. Currently, he is working on field 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 field provides valuable inspiration and is integral to main- taining his enthusiasm. He considers his notebooks, which he fills with written and sketch notes, as the source of his most valu- able references and inspirations. John encourages everyone interested in natural history to keep a notebook.

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