Table Of ContentORNITHOLOGY
in Laboratory and Field
Fifth Edition
Olin Sewall Pettingill, Jr.
Laboratory of Ornithology
Cornell University
Illustrated by
Walter! Breckenridge
University of Minnesota
Bell Museum of Natural History
/ft
Academic Press, Inc.
(Harcourt Brace Jovanovich, Publishers)
Orlando San Diego San Francisco New York
London Toronto Montreal Sydney Tokyo Säo Paulo
To the memory of
Alfred O. Gross
(1883-1970)
Professor of Biology at Bowdoin College,
preeminent ornithologist and inspiring teacher.
Copyright © 1985 by Academic Press, Inc.
All rights reserved.
No part of this publication may be reproduced or
transmitted in any form or by any means, electronic
or mechanical, including photocopy, recording, or
any information storage and retrieval system, without
permission in writing from the publisher.
Academic Press, Inc.
Orlando, Florida 32887
United Kingdom Edition Published by
Academic Press, Inc. (London) Ltd.
24/28 Oval Road, London NW17DX
ISBN: 0-12-552455-2
Library of Congress Catalog Card Number: 84-71089
Printed in the United States of America
Preface
Like its four predecessors, this fifth edition of Or- Daily field checklists (as many as there will be
nithology in Laboratory and Field is intended for field trips)
use at the college or university level and assumes
Loose-leaf pocket notebook, preferably with alu-
that students need have only a background knowl-
minum cover
edge of general biology. Although the book makes
no pretense of covering the realm of ornithology, it
A set of colored crayons
treats those major aspects of the science that can
be studied during a course in the regular academic
The institution should make available to students
year or in the summer.
the following:
There are twenty-two chapters, all more or less
independent units. Even though they are pre-
The American Ornithologists' Union's Check-list
sented in a fairly logical sequence, they can be taken
of North American Birds, Sixth Edition
up in almost any order, and some may be omitted
altogether without affecting the value of the others. Annotated checklist of birds in the region of the
The instructor may wish to select parts of the book study
without regard to sequence that will best supple-
ment class work or suit the season of the year during A world atlas
which the course is given.
Telescope (preferably the so-called "spotting
With few exceptions each chapter concludes with
scope") with either eyepiece or a zoom lens and a
references. While many references are cited in the
tripod
preceding text of each chapter, many more are in-
cluded to support or supplement the text, to give Portable tape recorder
views contrary to statements in the text, or to sug-
Compound microscope
gest further reading. If the titles are not cited from
the text or the titles themselves do not indicate the Meter stick
reasons for their inclusion, they are annotated par-
A Common Pigeon (Rock Dove) and a House
enthetically. Students should make a practice of
Sparrow, properly preserved but not plucked, for
perusing the references after reading the preceding
external study and dissection
text.
In order to use this book effectively, certain Several nestling House Sparrows preserved in
equipment and materials are desirable. The student spirit for study of feather tracts
should have for personal use:
Pigeon skeleton, mounted
Binocular
Pigeon skeleton, completely disarticulated
Field guide
Key to the bird species in the region where the Human skeleton, mounted, or a detailed chart of
study is made a human skeleton
IX
x / Preface
Human cervical vertebra (fifth) following: Figures 1-3, 5-6, and 24 by William
Montagna; Figures 4 and 38 by Ray S. Pierce; Fig-
Hyoid apparatus of a woodpecker
ures 8-11 by Robert Gillmor; Figures 12-23, 25-
A series of contour feathers illustrating special- 28, 34, and 36 by Robert B. Ewing; Figures 29,
ized feather types. Some of the more exotic types 30a, and 31 by Barbara Downs; Figure 33 by Sandra
may be obtained from zoos, which usually save such L.L. Gaunt; Figure 41 by Frank A. Pitelka (first
material for educational institutions. published in The American Midland Naturalist for
1941); Figures 51-53 by Sidney A. Gauthreaux, Jr.;
A semiplume, an adult down feather, a filoplume,
and Figure 54 by Helen S. Chapman.
a bristle, and a section of the vane of a contour
For the new chapter "Flight" in this edition, I
feather mounted on slides for microscopic study.
am grateful to the Laboratory of Ornithology at
Parts of feathers with different colors—red, or- Cornell University for granting me permission to
ange, yellow, black, gray, brown, iridescent, blue, use the drawings by Robert Gillmor and parts of
green, and white—mounted on slides for micro- the accompanying text from the Laboratory's Home
scopic study of color-producing elements Study Course in Bird Biology, Seminar IV.
I wish to thank the Bird Banding Laboratory at
A series of spread wings of a passerine species Patuxent, Maryland, for reviewing in Appendix A
(e.g., the House Sparrow or European Starling) to the procedures and regulations for obtaining per-
illustrate the progress of molt mits to capture, band, and mark wild birds, as well
as for providing addresses of state and provincial
A collection of bird skins representing all the
agencies to which inquiries and applications should
orders and families of North American birds and all
be made. I also wish to thank Janet G. Hinshaw,
species found in the region of the study. (If possible,
Librarian in the Josselyn Van Tyne Memorial Li-
the collection should be sufficiently comprehensive
brary at the University of Michigan, for help in
to show sex, age, and other constant differences in
checking on, and adding to, the titles of various
plumage; abnormal plumage coloration; color
ornithological journals in Appendix C.
phases; eclipse plumage; and plumage changes by
In the preparation of this edition I am deeply
wear and fading.) A transparent plastic tube
indebted to the following authorities in their re-
(capped) for each of the smaller skins that will be
spective fields of ornithology:
handled often is recommended in order to prevent
Sidney A. Gauthreaux, Jr., for writing the entire
damage. (These can be ordered from any biological
chapter "Migration."
supply company.)
Jack P. Hailman, for writing the entire chapter
A record player or tape recorder with reproduc- "Behavior" and for reviewing all the succeeding
tions of songs of species occurring regularly in the chapters on the breeding biology of birds.
region of the study Peter Stettenheim, for again giving detailed at-
tention to the chapter "Feathers and Feather
The prefaces to the previous editions of this book Tracts," further clarifying or amplifying many as-
tell the story of how it evolved, starting in 1939. In pects of feather growth, structure, coloration, and
these prefaces I have acknowledged ornithological function.
sources for much of the text. I also acknowledged Andrew J. Berger, for reviewing and updating
my many colleagues, former students, numerous the "Muscular System," and for taking the time to
friends, and my late wife, Eleanor, who contributed go over all parts of the book pertaining to internal
in various ways to the production of all four previous anatomy and physiology.
editions. Without the advice or assistance of these Abbot S. Gaunt, for describing the anatomy of
generous people I would not have had the frame- the syrinx and its function.
work and much of the authoritative information in Kenneth C. Parkes, for reviewing and, in some
this edition. instances, revising or rewriting parts of the chapters
The cover illustration is by Walter J. Brecken- "Systematics and Taxonomy" and "Plumages and
ridge and all the illustrations in the text, if not Plumage Coloration"; also for overseeing the exact-
accompanied by credit lines, are his except the ing work of Jay Loughlin, Collection Manager at
Preface I xi
the Carnegie Museum, in updating the synopses of populations and giving instructions for a new
North American orders and families of birds and method of measuring populations.
reconstructing the keys to the same orders and Alan Feduccia, for his critical reading of the an-
families in accordance with the sixth edition (1983) cestry and evolution of birds in the last chapter of
of The A Ό.Ό. Check-list of North American Birds. this book.
Donald E. Kroodsma, for greatly expanding the Robert A. McCabe and Ray B. Owen, Jr., for
chapter "Song" and giving the benefit of his ex- reading and commenting in Appendix A on field
pertise in recording bird vocalizations under "Or- techniques applied to capturing, marking, and fol-
nithological Field Methods" in Appendix A. lowing wild birds.
Stephen I. Rothstein, for significantly extending Finally, I thank my long-time friend, Edward F.
the sections on brood parasitism in the chapters Dana, with his considerable editorial experience,
"Eggs, Egg-laying, and Incubation" and "Young for reading much of the text of this edition when it
and Their Development." was in galley proof.
John T. Emlen, for worthy information on bird Olin Sewall Pettingill, Jr.
Birds and Ornitl·
An Introduction
Birds among all animals offer the most favorable take flight, to stay aloft, and to reach its destination
combination of attributes for scientific study. They under the most adverse conditions.
are numerous, abundantly diversified in form, and
easily observed. They are highly organized and re-
Achievements for Flight
sponsive with sensory capacities similar to mans
and therefore understandable. Pleasing in colors Several achievements have contributed to the
and movements, they are also, with few exceptions, bird's mastery of the air.
inoffensive in their habits and incapable of physi-
cally harming the investigator. Many adapt readily Lightness Achieved by a covering of feathers—
to experimentation. Little wonder that ornithology, "the strongest materials for their size and weight
the science of birds, boasts so many practitioners, known"—instead of a thick skin; by the loss of teeth
and in turn contributes so significantly to modern and the heavy jaws to support them; by a reduction
concepts of evolution, speciation, behavior, and of the skeleton and by the hollowing, thinning, and
ecology. flattening of the remaining bones; by a radical short-
ening of the intestine and the elimination of the
urinary bladder; and by air spaces in the bones,
Birds Defined body cavity, and elsewhere.
Birds are unique among all animals in being feath- Streamlining Also achieved by the feathers, over-
ered. Like mammals, they too are warm-blooded, lapping and smoothing the angular, air-resistant
or homeothermous, (capable of regulating their surfaces and providing bays, wherein the feet may
body temperature). And like most of their verte- be withdrawn.
brate associates, excepting most mammals and a
few others, they lay eggs. Centralization and Balance Achieved by posi-
Animals move from place to place by running, tioning all locomotor muscles toward the body's
hopping, walking, crawling, swimming, gliding, center of gravity—leaving the wings, like puppets,
and flying. Among birds, flight is the principal controllable by tendinous strings; and by position-
means of locomotion, even though some forms— ing the gizzard, the avian substitute for teeth, and
for example, ostriches, kiwis, and penguins—in the other heavy abdominal organs in the center of the
course of evolution have lost their ability to fly. body.
Therefore, one recognizes birds as birds because
they are formed to fly. Maximum Power Achieved by the combination of
The modern bird, like an airplane, is structurally an exceptionally high, steady body temperature for
and functionally efficient. A bird must be able to aerial maneuvers in all extremes of climate and
1
2 / Birds and Ornithology: An Introduction
weather; by feathers, which aid in conserving the attain. They cannot be as small or as large as many
heat; by increased heart rate, more rapid circulation other animals. Because they have a high rate of
of the blood, and greater oxygen-carrying capacity metabolism for supporting a high body temperature
of the blood stream; by a unique respiratory system, and flight movements, birds need sufficient food to
which permits a double tide of fresh air over the maintain this rate and at the same time compensate
lung surfaces, synchronizes breathing movements for heat loss from body surfaces.
with flight movements, cools the body internally, Theoretically, the smaller the bird, the greater
and eliminates excess fluids; and by a highly selec- is its relative body surface in relation to weight and
tive diet of energy-producing foods, which contain the greater its heat loss. Consequently, the smaller
few indigestible substances to cause excess weight. the bird, the more it must eat in proportion to size.
Again, theoretically, a bird smaller than kinglets
Visual Acuity and Rapid Control Achieved by and chickadees would have to eat all the time, night
large eyes with a wide visual field and remarkable and day. Hummingbirds exist, small as they are,
distance determination, and by a brain whose because they lower their body temperature—that
greatly enlarged visual and locomotor centers are is, become torpid—at night or at other times when
capable of recording and transmitting nerve impul- they cannot eat. Thus they conserve energy.
ses with the reactions of a seasoned pilot. The larger the bird, the faster it must fly to stay
airborne. It needs bigger flight muscles for greater
speed. This, in turn, means greater weight because
Range in Size
flight muscles are heavy.
Birds range widely in size. The Ostrich (Struthio The larger birds have attained their size while
camelus), standing between 8 and 9 feet tall (2.44 retaining their ability to fly by developing a de-
and 2.74 m) and weighing nearly 350 pounds (159 pendence on air currents. Albatrosses and condors
kg), is the largest. But it is, of course, flightless. practically require winds and updrafts in order to
Among the largest flying birds are the Royal and fly at all.
Wandering Albatrosses (Diomedea epomophora
and D. exulans) and the Andean and California Con-
dors (Vultur gryphus and Gymnogyps californi-
Ornithology Defined
anus) with^ingspans approximating 10 feet (3 m).
The Marabou Stork (Leptoptilos crumeniferus) may
be the largest flying bird, if, as reported, its wing- Ornithology, simply defined, is the science of birds.
span measures over 12 feet (3.7 m). For a descriptive definition, there is none more
The smallest birds include numerous species of suitable than the one written by Elliott Coues, the
hummingbirds, the extreme being the Cuban Bee perceptive American ornithologist, nearly a cen-
Hummingbird (Mellisuga helenae) that measures tury ago:
2.25 inches (5.72 cm) from bill-tip to tail-tip and
Ornithology consists in the rational arrangement and
weighs less than 2 grams. Fourteen Bee Hum- exposition of all that is known of birds, and the logical
mingbirds would weigh no more than an ounce inference of much that is not known. Ornithology treats
(28.35 g). of the physical structure, physiological functions, and
Within a species there is often sexual difference mental attributes of birds; of their habits and manners;
in size, the males averaging slightly larger. In some of their geographical distribution and geological
species sexual dimorphism is very marked with the succession; of their probable ancestry; of their every
relation to one another and to all other animals, in-
male slightly more than twice as large, as in the
cluding man. [In Key to North American Birds, 5e
Wild Turkey (Meleagris gallopavo) and in the larg-
(Boston: Dana Estes and Co., 1903), p. 58.]
est of all grouse, the Capercaillie (Tetrao urogallus),
or with the female a third larger, as in the Sharp- One must study ornithology in both laboratory
shinned and Cooper's Hawks (Accipiter striatus and and field because a knowledge of birds "in the hand"
A. cooperii). is incomplete without a knowledge of birds "in the
There are limits to the size that flying birds may wild," and vice versa.
Ornithology Defined I 3
Form, Structure, and Physiology shape, plumages and coloration, breeding habits,
seasonal movements, and general behavior. Or, to
Basic to the study of ornithology is an introduction
put it another way, thanks to adaptive radiation
to the form, structure, and physiology of birds. This
operating so vigorously in the descent of birds,
the student can best accomplish by making direct
there are some 9,000 different species today.
observations on the physical make-up of a "gener-
In studying birds one naturally thinks of them in
alized" bird, such as the Rock Dove or Common
terms of species. Therefore, the logical sequel to a
Pigeon (Columba livia), and by learning from a text
knowledge of their form, structure, and physiology
book the role of each organ system in the bird's way
is an acquaintance with the many different species
of life. Attention must be centered on those features
in the student's immediate area. This requires un-
that will particularly enhance an appreciation of
derstanding the concept of species and speciation
birds as biological entities. Frequently, certain fea-
and the methods of classifying, naming, and
tures must be compared to their homologues in
identifying.
man, thereby making them more understandable.
Gaining a thorough acquaintance with the 150 to
The logical sequence to such an introduction is,
300 species regularly occurring in the average study
first, the identification of the different parts of the
area of temperate North America demands a knowl-
bird's topography, followed by a study of the bird's
edge of the taxonomic characters and other means
feather covering—how the feathers are structured
of recognizing species in both laboratory and field,
and variously modified, how they develop, how
together with an understanding of changes in plu-
they are colored, and how they are arranged on the
mage and plumage coloration among different
body. A detailed knowledge of these exterior fea-
species.
tures is indispensable not only in describing birds
The identification of species is not an end in itself
and their actions but also in accounting for many of
but a stepping stone to investigations of many as-
their adaptations.
pects of bird life, or of biological problems in which
With this knowledge, the student is then pre-
birds play a role. Some students find speciation per
pared to investigate the internal organ systems.
se a challenging field since there is still much to be
Avian anatomy and physiology offer many oppor-
learned about the origin, status, and interrelation-
tunities for research. Indeed, an increasing number
ships of species.
of ornithologists specialize in one or both of these
fields, dealing particularly with the adaptive and
comparative aspects among different species of Distribution
birds.
Although most modern species of birds can fly and
thus can rove the earth, each species is confined to
Species and Speciation
a particular geographical range, which may be from
Although uniformly specialized for flight, birds several hundred acres, as on a sea island, to one or
have nonetheless changed widely in form and action more continents in size.
in order to live in particular environments. The ranges of species overlap so that in any one
Consider, for example, the adaptations for loco- area there is an aggregation of species—an avifauna.
motion and feeding. Some species customarily fly Because the ranges of species are rarely or never
swiftly; others fly slowly. Some hover; others soar. identical, avifaunas vary markedly. Students over
Some swim and dive; others wade. Some walk or the years have given attention to the composition,
hop; others climb. To get food, some species probe comparison, and origin of avifaunas, yet there is
in the soil, others dabble in shallow water, scratch much about them that remains to be investigated.
the ground, chisel holes in trees, make flying sor- Geographical ranges are unstable due partly to
ties, or hunt for prey in any number of different the tendency among species to invade new areas.
ways. Cyclonic storms may help or hasten resettlement
These adaptations and others, always in complex by moving individuals to a different place, where
combination, account for the different shapes of they survive and reproduce if the environment suits
wings, tails, bills, and feet and differences in body them. Man has a part in it, too, when, for example,
4 / Birds and Ornithology: An Introduction
he transports birds on his ships. House Sparrows Rarely do they find one species throughout a com-
(Passer domesticus) reached the Falkland Islands in munity, even though it may be characteristic ofthat
the South Atlantic on ships that first stopped at environment. As a rule, it occupies merely a niche
Montevideo, Uruguay, where the birds, attracted and is adjusted to this position in structure, func-
to sheep-pens on deck, came aboard and remained tion, and behavior as no other species in the same
until the ships reached the islands. community. In the forest community, for example,
Any student, after having observed birds in a the Red-eyed Vireo occupies a treetop niche and is
given area for a few years, is certain to note shifts adjusted to this position in structure, function, and
in ranges and ponder the reasons. Modern orni- behavior just as the Ovenbird (Seiurus aurocapil-
thologists pay considerable attention to local distri- lus) occupies the forest floor. It would be unusual
bution as the abundant literature on the subject to see the Red-eyed Vireo on the ground or the
clearly indicates. Ovenbird in the treetops. While a species may ap-
Within its geographical range a species, if nor- pear to share its niche with other species, not one
mally migratory, is seasonally distributed, appear- behaves exactly as another does or requires the
ing in one part of its range in one season, in another same food and the same nesting site.
part in another season. Bird species are of greater variety and density in
Within its geographical range a species is also ecotones than in the pure communities that border
ecologically distributed. It usually occupies a par- them. This phenomenon, called edge effect, is im-
ticular environment or habitat and shares this hab- portant to anyone wishing to see larger numbers of
itat with other organisms—plant and animal—all of birds.
which are adapted to the prevailing conditions of Edge effect results in a greater variety of vege-
soil, air temperature, moisture, and light. All the tation—grasses, shrubs, and trees—providing a
organisms in a given habitat collectively comprise greater variety of food and cover for birds. For
a biotic community, since they show relationships example, ecotones where field and forest merge
to one another. have the plants characteristic of both field and forest
When any two communities meet, more often and many additional shrubs. Thus they bring to-
than not, there is an area of mixture and overlap, gether birds of both field and forest and also attract
or ecotone, in which the birds and other living species that require either shrublands or a combi-
forms characteristic of these communities are in- nation of trees, shrubs, and grasses.
termixed and in which are additional forms that, Some bird species are adapted so strongly to a
preferring this ecotone, seldom occur elsewhere. special niche that they cannot live in a different
Students soon become aware of the importance situation. If an element in the niche on which they
of habitat or community in accounting for the pres- depend is destroyed or seriously altered, they are
ence or absence of species and, before long, learn more likely to disappear than to make an adjust-
to associate different species with particular envi- ment. The Snail Kite (Rostrhamus sociabilis) prob-
ronments—the Red-eyed Vireo (Vireo olivaceus) ably would disappear in Florida were disaster to
with the deciduous forest, the Horned Lark (Ere- befall the big freshwater snail, Pomacea palludosa,
mophila alpestris) with the short-grass prairie, and on which it feeds exclusively. It is likely that the
the Verdin (Auriparus flaviceps) with the scrub Kirtland's Warbler (Dendroica kirtlandii) would
desert. When students travel northward on the con- disappear in northern Lower Michigan, where it
tinent of North America or climb a high mountain, breeds exclusively, if there were no more jack pines
they expect a sequence of species as they pass 6 to 18 feet high (2 m to 6 m) under which it almost
through one environment after another—the Olive- invariably nests.
sided Flycatcher (Contopus borealis) in the con- A good many bird species, on the other hand,
iferous forest, the White-crowned Sparrow (Zono- are much more adaptable. Sometimes they are so
trichia leucophrys) at the timberline ecotone, and widely tolerant of different situations that their pre-
ptarmigan (Lagopus spp.) on the tundra. cise niches are unrecognizable. The Blue Jay (Cyan-
At the same time students become conscious of ocitta cristata), Black-capped Chickadee (Parus
several significant aspects of ecological distribution. atricapillus), and Cedar Waxwing (Bombycilla ced-
Ornithology Defined I 5
rorum) are so adaptable that one may find them Many mating displays are actually derived from
almost everywhere in wooded areas through their such maintenance activities as preening or scratch-
ranges. ing; or from displacement activities—for example,
The species that restrict themselves to narrowly when a bird breaks off fighting and pecks at some
prescribed niches generally have small populations object; redirected activities—when a bird redirects
within correspondingly small ranges. The species its attack to an object other than one which elicited
tolerant of environmental changes and variations the response; and intention movements—when a
are mainly the inhabitants of the ecotones; they bird makes a move to fly but fails to do so, thereby
have large populations and often range widely. performing an incomplete act.
The underlying factors accounting for the ecolog- Inherited behavior predetermines the extent to
ical distribution of many species still remain to be which learned behavior may develop. Learned be-
determined. Here is a study with a degree of ur- havior is actually adaptive behavior resulting from
gency. As man steadily destroys the natural envi- experience. A bird inherits the ability to fly, yet it
ronments, an understanding of a species' ecological must learn by experience to take off into the wind
requirements is the first step in preventing its de- rather than with it and to choose the perch that will
crease. The next step is to see that its requirements best accommodate its feet. This is called learning
are maintained through intensive management and by trial and error. Other forms of learning are by
conservation practices. habituation and by imprinting. A few birds show
ability to learn by insight. The different methods of
learning among birds demand much more research.
Behavior
Investigators often give considerable attention to
The behavior of birds attracts scores of investiga- social behavior since most birds are by nature gre-
tors. Birds are ideal animals for behavioral studies. garious and have consequently developed many
Each species has an impressive repertoire of innate kinds of interactions related to attack, escape, de-
behaviors and, at the same time, its ability to learn fense, flocking, and reproduction. Although the lit-
compares favorably with that of most mammals. erature on social behavior in birds is already
Thanks to a rich variety of bird species, each with enormous, the subject is still a fertile field for study.
a different mode of life, investigators have avail-
able for study a correspondingly rich variety of
Migration
behaviors.
An understanding of the principles of bird be- No aspect of bird life has so excited man's interest
havior is essential for any beginning student, help- down through the centuries as the withdrawal of
ing as it does to explain the basis of many avian birds from an area in the colder seasons and the
activities. Even more important, an understanding return to the same area when the seasons become
of bird behavior illuminates many of the basic etho- warmer. In spite of a great store of knowledge on
logical principles applied to human life. Modern the initiation and procedure of migration among
psychologists are now paying attention to such phe- modern birds, the question of how and when mi-
nomena as individual distance and dominance re- gration originated still remains speculative—an
lationships (first noted in birds!) that are so evident ever-present challenge to one's thinking.
in urban societies. Continued, in-depth studies of Experimental studies started over 50 years ago
avian behavior will, almost certainly, further demonstrate that a specific day length in the spring
sharpen man's perception of his own social stimulates the activity of a bird's endocrine glands,
problems. and this stimulation brings the bird into a migratory
The procedure in the study of behavior is to state. Some external factor then releases migratory
identify, describe, and name the behaviors of a behavior. In the fall, with a regression of endocrine
species and then to determine what each behavior activity, the bird reaches another migratory state
accomplishes, its significance to the species' sur- ready for triggering by an outside cause.
vival, its causes, how it has evolved, and whether The present wealth of information on the process
it is innate, learned, or both innate and learned. of migration—starting and stopping times, rate,
