The Real Roadrunner (Animal Natural History)

The Real Roadrunner

ANIMAL NATURAL HISTORY SERIES Victor H. Hutchison, General Editor

The Real Roadrunner MARTHA ANNE MAXON

UNIVERSITY OF OKLAHOMA PRESS : NORMAN

Library of Congress Cataloging-in-Publication Data Maxon, Martha Anne, 1941– The real roadrunner / Martha Anne Maxon. p. cm. — (Animal natural history series) Includes bibliographical references (p. ). ISBN 0-8061-3676-6 (alk. paper) 1. Roadrunner. I. Title. II. Series. QL696.C83M39 2005 598.7'4—dc 22 2005041716 The Real Roadrunner is Volume 9 in the Animal Natural History Series. The paper in this book meets the guidelines for permanance and durability of the Committee on Production Guidelines for Book Longevity of the Council on Library Resources, Inc. ∞ Copyright © 2005 by the University of Oklahoma Press, Norman, Publishing Division of the University. All rights reserved. Manufactured in the U.S.A. 1 2 3 4 5 6 7 8 9 10

To the memory of my father, Byron Martin White, DVM, who loved wild things and taught me to love them, and to the memory of Dr. George Miksch Sutton, who loved the roadrunner and taught me to love it.

Contents

List of Illustrations

ix

Preface

xi

Acknowledgments

xiii

Chapter 1. Roadrunner Basics

3

Chapter 2. Food and Foraging

17

Chapter 3. Daily Survival

29

Chapter 4. Breeding Characteristics and Courtship

37

Chapter 5. Getting Together, Nest Building, and Incubating

49

Chapter 6. Caring for Nestlings and Fledglings

59

Chapter 7. Development of the Young

67

Chapter 8. Population and Range

77

Chapter 9. A Bird of the People

91

Bibliography

107

Index

117

Illustrations

Plates Following page 50 1. Adult roadrunner in Chihuahuan Desert of West Texas 2. Male roadrunner with crest erect and colored skin patch displayed 3. Male roadrunner with crest partially erect and colored skin patch displayed 4. Recently fledged roadrunner, about twenty-five days old 5. Roadrunner starting to swallow Texas horned lizard 6. Roadrunner swallowing Texas horned lizard 7. Juvenile roadrunner absorbing heat from the sun on the bare black patch on its back 8. Roadrunner sunning during the heat of the day 9. Captive male roadrunner holding Chihuahuan spotted whiptail lizard during the tail-wagging display of courtship 10. Mating of roadrunners 11. Roadrunner nest with five eggs under attack by bull snake 12. Roadrunner nest with ten eggs 13. Female roadrunner incubating ten eggs 14. Coiled western diamondback rattlesnake 15. Nesting adult roadrunner attacking a taxidermic roadrunner mount placed near its nest 16. Roadrunner nest with four chicks, four days old and younger 17. Twenty-five-day-old juvenile in immobile, erect posture 18. Month-old juvenile begging for food from its parent 19. Head of four-to-five-day-old chick

Illustrations

x

20. Foot and wing of four-to-five-day-old chick 21. Fifteen-day-old roadrunner nestling 22. Twenty-three-day-old juvenile 23. Two-month-old juvenile 24. Seventy-five-day-old juvenile

Figures 1. Yellow-billed cuckoo 2. Zygodactyl foot of roadrunner 3. Rufous-vented ground-cuckoo 4. Wing and leg of roadrunner 5. Variations of crest erection and color patch display 6. Roadrunner flashing its wings to flush prey 7. Head movements of a six-note coo call 8. Prance display of a male 9. Tail-wag display 10. Female taking lizard from male at end of mating 11. Mating with no food exchange 12. Mutual flick display by both sexes after mating 13. Female giving the whine call from a perch 14. Thirty-five-day-old juvenile crouching after parental attack 15. Locations by county of roadrunner records from mid-1800s to present time 16. Range expansion of roadrunner from 1900 to 1975. 17. Roadrunner track 18. Petroglyph of mountain lion track and roadrunner track

5 6 6 10 12 23 41 43 44 45 45 46 47 64 78 82 94 96

Tables 1. General range and habitat of the roadrunner and its ground-cuckoo relatives 2. Breeding-season stages and approximate durations at a roadrunner nest 3. Development of nestling roadrunners from hatching to twenty days old 4. Development of fledgling roadrunners from twenty-one days old until first fall season 5. Common names for the roadrunner

8 38 72 73 103

Preface

A

brief glimpse of the roadrunner as it darts across the road is enough for many birdwatchers. Some of them, however, would like to know more about the roadrunner’s daily life. In my years of studying this bird, I found that with patience and persistence a person can become privy to many of its secrets. As Dr. George Miksch Sutton, who first inspired me to study the roadrunner, so aptly put it: “He who knows the roadrunner, he who has measured the breadth and the depth of this unique bird personality, has lived with him—not for an hour or so, not for a day, but week after week after week.” In this book, I share the roadrunner secrets I learned through spending many breeding seasons with them. I hope that the reader is able to experience vicariously the wonder I have experienced by observing this bird in the wild, which has been a major highlight of my life. I also hope that this book will stimulate further research into the many aspects of the roadrunner’s life that still hold secrets. A major flaw in our knowledge is that all of the field studies of roadrunners have been done in southwestern desert scrub or brush habitats, to the neglect of its natural history in the eastern and northern parts of its range. Throughout the book, I point out unanswered questions and suggest topics for further research. I hope that some of you may help answer them someday. Even if all the answers don’t come easily, the time spent with the roadrunner will be well worth it. This book is intended to acquaint the reader with the daily life of the roadrunner and with the amazing adaptability that has made it a survivor through the centuries. The book begins with some basics about the bird—its ancestry, anatomy, habitat, and range. Then we will look at how the roadrunner meets its basic survival needs of obtaining food, withstanding the elements, and avoiding predators. The heart of the book covers the breeding behavior of

xii

Preface

the roadrunner, which occupies its time for three-fourths of the year and illustrates the adaptable nature of this species. Understanding the natural history of the roadrunner sets the stage for exploring the reasons for its ability to expand its range five hundred miles eastward from its desert home into the heartland of the United States during the last century. Besides the bird’s adaptability, human changes to the landscape may have helped in its eastward trek. Barriers to its further eastward and northern expansion may exist, however, and we will examine them to round out our understanding of the roadrunner’s natural history. Because this bird has been so intimately connected with various cultures for many centuries and is popular in modern culture, I have included a final chapter on the colorful folklore and tales about the roadrunner. The roadrunner is one of the icons of the Southwest and one of the most loved birds of our country. In this book, for ease of reading, I use the term ‘roadrunner’ rather than its official common name, “greater roadrunner.” For those who are interested, the scientific names of plants and animals mentioned in the text are placed in the index, under the common name of each species. The sources I have used appear in the bibliography. Approximately two-thirds of these are cited in the text. The remainder provided background information, and I have included them in the interest of completeness.

Acknowledgments

long the way, I have been helped by many wonderful people too numerous to name. But I will attempt to acknowledge as many as I can.

A

Lawrence E. Maxon, my beloved husband, who never ceased to be my support and anchor throughout the writing of this book Dr. Byron Martin White and Ione Bullard White, my late father and mother, who helped me watch roadrunners and find nests and gave me a childhood full of the wonders of nature Byron White, my brother, who faithfully read most of this manuscript and provided comments and constant encouragement Dr. George Miksch Sutton (deceased), University of Oklahoma, who first taught me about the wonders of roadrunner-watching and shared many of his insights and questions about the bird, as well as his experiences in raising pet roadrunners as a young boy Dr. Frederick R. Gehlbach, Baylor University, who gave me an ecological viewpoint Dr. Charles C. Carpenter, University of Oklahoma, who taught me the principles of animal behavior, field observation, and time-motion analysis of behavior Dr. Victor Hutchison, general editor of the University of Oklahoma Press’s Animal Natural History Series, who provided editorial guidance Dr. Gary Schnell, University of Oklahoma, who provided several Oklahoma references Dr. Matthew C. Perry, Patuxent Wildlife Research Center, who spent time in a musty government storage room digging out the old U.S. Biological Survey food habits cards for the roadrunner and supplied them to me

xiv

Acknowledgments

Dr. Wade C. Sherbrooke, American Museum of Natural History, Southwestern Research Station, Portal, Arizona, who discussed his horned lizard and roadrunner research with me Polly Schaafsma, anthropologist and artist of Santa Fe, New Mexico, who took time to discuss American Indian petroglyphs and folklore with me Dr. Elizabeth Brandt, Arizona State University, who discussed American Indian folklore with me and directed me to major literature sources Barbie Coselman, who thoughtfully read this manuscript from the point of view of a layperson and a top-notch secretary and gave me invaluable suggestions on content and grammar Terry van de Walle, who enthusiastically conducted the records search for past and current roadrunner distribution Dr. Janice M. Hughes, who shared her insights on cuckoo phylogeny and her observations and enthusiasm for the roadrunner James D. Lowe, Cornell Laboratory of Ornithology, Cornell University, who provided the nest record card data from the lab’s Bird Population Studies database Kathy Klimkiewicz, Bird Banding Laboratory, Patuxent Wildlife Research Center, Laurel, Maryland, who provided bird-banding data Jan Bird, Kimberly Gilbert, Willie Graham, Jewel Schultheis, and Pat and Dick Weekly, all dear friends whose friendship and prayers saw me through to the end Jacque Staston, graphic artist, who digitally edited plates 1–4, 10, 11, 16, and 21 in this book and shares my enthusiasm for roadrunners Ione Prange and Janice Holmes, librarians who tracked down many of the reference articles used for this book Staff of the University of Oklahoma Press who guided this manuscript to completion, including Kimberly Wiar, Jean Hurtado, Karen C. Wieder, and Marian Stewart Rosemary Wetherold, another roadrunner fan, who did a magnificent job as copy editor of the book The many volunteers who have contributed to our knowledge of roadrunner nesting habits by participating in the North American Nest Record Card Program, and especially to Mr. and Mrs. Orphus C. Bone, who covered five counties in the South Texas brushlands between 1967 and 1974 and provided almost a third of the roadrunner records The following providers of financial support for my roadrunner field research over the years: a National Park Service Research Grant, an NSF Grant-in-Aid at the University of Oklahoma Biological Station, and the National Geographic Society

Illustration Credits Curtis Schaafsma (provided by Polly Schaafsma): figure 18. Dr. Wade Sherbrooke: plates 5, 6, and 9. All other photographs are my own. All drawings are by Mark Hughes, Hudson, Quebec, Canada, with the exception of figures 5, 15, and 16, which are my own.

The Real Roadrunner

1 Roadrunner Basics

riting this book has brought back wonderful memories of time I spent with the roadrunner in Texas and New Mexico. Looking back, I realize how privileged I have been to observe the life of this amazing bird in its natural habitat. I also raised several of these birds in captivity as part of my study of their courtship behavior for my doctoral dissertation. Altogether, I spent six breeding seasons with these birds, involving thousands of hours of observation and many still photographs and movie films of behavior and tape recordings of calls. I feel privileged to share what I and others have learned about this intriguing bird with the readers of this book. My romance with the roadrunner began when I was a child growing up in the brush country of South Texas. I was fortunate to have a father who loved the animals and land of this area of Texas. Our family’s outings were spent learning about the wild things around us. One of the creatures that fascinated me the most was the roadrunner, whose charisma entrances those fortunate enough to encounter it. It is hard to pin down exactly what facets of this complex bird grab one’s attention, but no one who has spent time with this bird remains dispassionate about it, whether scientist or layperson. My hope is that this book will acquaint the reader with the life of the roadrunner and stimulate interest in helping uncover its many secrets that still elude us. Although the roadrunner is an icon of the Southwest and beloved by people throughout the world, it has been one of the most misunderstood of birds. This misunderstanding has come about not only because ideas about the roadrunner are colored by centuries of folklore

W

4

The Real Roadrunner

and the twentieth-century animated character but also because little knowledge existed about the real bird until relatively recently. Roadrunner research of the last fifty years has revealed a real creature every bit as fascinating as the legend and cartoon, if not more so. As the first step on our journey to know the real roadrunner, we begin with its discovery by Euro-American people. An anonymous Franciscan priest wrote the first known description of the roadrunner in 1790. His account appeared in a Spanish manuscript containing memories of California’s natural history: “The Churca is a kind of pheasant which has a long bill, dark plumage, a handsome tail and four feet. It has these latter facing outward in such fashion that when it runs it leaves the track of two feet going forward and two going backward” (quoted in Coues 1900, 70; note that the “four feet” could be a mistranslation of the Spanish). Although an astute observer of the roadrunner’s anatomy, this priest made the same mistake that many casual observers of this bird do: grouping it with the pheasant. The next report comes from famous explorer Lieutenant Zebulon Montgomery Pike, who caught a roadrunner in 1806 in Chaffee County, Colorado (A. M. Bailey and Niedrach 1965). However, it wasn’t until 1829 that French naturalist and surgeon RenéPrimevère Lesson provided the first scientific description of the bird from a San Diego County, California, specimen. He gave it the scientific name Saurothera californiana, correctly identifying it as a member of the cuckoo family. Its placement in the genus Saurothera allied the roadrunner with several distantly related New World cuckoos, the lizard-cuckoos. In 1831 the roadrunner was placed in the new genus Geococcyx, along with a newly discovered species, the lesser roadrunner, Geococcyx velox. Between then and 1858 the roadrunner was known by various scientific names, including Geococcyx variegata, G. mexicanus, G. viaticus, G. marginatus, and G. californicus (Ridgway 1916). Geococcyx californianus, the current scientific name for the roadrunner, was established in 1858 by Spencer Fullerton Baird, assistant secretary of the Smithsonian Institution (and later its director). The genus name means “earth [geo] cuckoo [coccyx].” The type specimen came from California; hence the species name californianus. The official common name for the species is “greater roadrunner,” to distinguish it from the lesser roadrunner. In this book, the term “roadrunner” is used to refer to the greater roadrunner. When the lesser roadrunner is mentioned, its full common name is used.

Roadrunner Roots As a background for the natural history of the roadrunner, we will look at it in the context of its cuckoo family. First impressions can deceive observers into thinking that the roadrunner is a fowl, that is, a cousin to the quail, the pheasant, and the chicken. But its ground-dwelling habit and drab appearance are misleading, for the roadrunner is a genuine member of the cuckoo clan. However, this earthbound bird is a nonconformist, standing in stark contrast to its aerial, tree-dwelling cuckoo relatives.

Roadrunner Basics

5

Figure. 1. Yellow-billed cuckoo.

To most, the word “cuckoo” brings to mind either a cuckoo clock or a crazy idea or person. In fact, the cuckoo-clock bird is patterned after the European cuckoo. The cuckoo is named for the cuc-cooing courtship call of the male European cuckoo. Because this bird gives his cry repeatedly, even late into the night, “cuckoo” came to describe something or someone dull, foolish, or crazy (Amon 1978). The order Cuculiformes, to which both the roadrunner and the European cuckoo belong, is a varied assortment of species. Besides the worldwide-occurring cuckoos, this group includes the coucals of Africa and Asia, the couas of Madagascar, and the anis and groundcuckoos of the Americas. This is the only order of birds with living species of tree dwellers that are strong fliers and ground dwellers that are weak fliers. The other ground-dwelling birds, such as the ostrich and the emu, do not have living tree-dwelling relatives. The grounddwelling quail, pheasant, and grouse are good fliers, unlike the ground-dwelling cuckoos. Close comparison of roadrunner features with those of other cuckoos, such as the yellowbilled cuckoo, reveals many similarities. Can you see resemblances between the roadrunner in plates 1–4 and the yellow-billed cuckoo in figure 1? Cuckoo species, whether treedwelling or ground-dwelling, share a set of physical characteristics, including the following: Long, stout, downward-curving bill Long, slender tail with white spots on tips of tail feathers

Figure 2. Zygodactyl foot of roadrunner.

Figure 3. Rufous-vented ground-cuckoo.

Roadrunner Basics

7

Zygodactyl foot, which has two toes pointing forward and two backward (fig. 2) Naked, two-lobed oil gland at the tip of the tail bone Bare patch of skin, usually colored, in the area behind the eye, having the effect of emphasizing the eye Rather drab plumage, often streaked Eyelashes Look-alike sexes Hatchlings naked or with sparse, hairlike strands on the body Thirty-six genera, containing 139 cuckoo species, are found throughout the warm and temperate areas of the world. A special aspect of cuckoo biology that has received attention from research biologists is brood parasitism, which occurs when the cuckoo species lays its eggs in the nest of another bird species. The host bird incubates the eggs and cares for the cuckoo young, usually at the expense of its own eggs or offspring. Often the color and pattern of the cuckoo egg match those of the host bird’s eggs. About 40 percent of cuckoo species practice brood parasitism. Roadrunners are part of the Neomorphinae, the ground-cuckoos/roadrunner subfamily of cuckoos. Cuckoo species in this subfamily occur only in the Americas. These species share features related to their ground-dwelling habit—long, strong legs and short wings. The long tail of the tree-dwelling cuckoos provides balance for perching and flying, and that of the ground-cuckoos provides stability as they walk or run. The species considered most closely related to the roadrunner are in the genus Neomorphus. Figure 3 shows one of these species, the rufous-vented ground-cuckoo of Central and South America. Table 1 provides general information about the Neomorphinae. The earliest known ground-dwelling cuckoo fossil is that of Cursoricoccyx geraldinae from a Colorado site. It is placed in the Neomorphinae group and may be closely related to today’s roadrunner species (Martin and Mengel 1984). Among the Neomorphinae, the roadrunner is the only species whose natural history is well known. The secretive habits, remote locations, and densely vegetated tropical or subtropical habitats of most of the ground-cuckoos make them hard to observe. Also, little information exists on the lesser roadrunner, despite its more accessible range and habitat. Information on a few ground-cuckoos appears in several places in this book for comparison with the roadrunner’s natural history. Another roadrunner, Conkling’s roadrunner, once roamed the Southwest during the late Pleistocene. Bone measurements indicate that this bird could have been 10 to 20 percent larger than the modern roadrunner. Paleontologists now consider it to be a subspecies (Geococcyx californianus conklingi) of the fossil, modern-sized form, Geococcyx californianus californianus, which occurred at that time as well (Harris and Crews 1983; Carpenter and Mead 2003). It is thought that there was some geographic separation of the two subspecies. To date, the majority of G. c. californianus fossils have come from Southern California, especially along the western coast, whereas the fossils of Conkling’s roadrunner have come from

Table 1 General range and habitat of the roadrunner and its ground-cuckoo relatives Scientific name Common name Length

General range

General habitat

Tapera naevia Striped cuckoo 11–12"

Mexico to Argentina

Dense thickets, brushy fields

Morococcyx erythropygus Lesser groud-cuckoo 10"

Mexico, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica

Arid woodlands, thickets, shrubby fields

Dromococcyx phasianellus Pheasant cuckoo 15–16"

Brazil, Peru, Colombia, Venezuela, Dense forest undergrowth, thickets, Paraguay humid forest edges

Dromococcyx pavoninus Pavonine cuckoo 10"

Brazil, Peru, Bolivia, Venezuela, Paraguay

Tropical rain forests

Geococcyx californianus Greater roadrunner 22–24"

United States, Mexico

Desert scrub, shrubby fields, open woods, riparian woods

Geococcyx velox Lesser roadrunner 20–22"

Mexico, Guatemala, El Salvador, Honduras, Nicaragua

Arid scrub, thickets, open woods

Neomorphus geoffroyi Rufous-vented ground-cuckoo 19–20"

Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Bolivia, Brazil

Tropical rain forest

Neomorphus squamiger Scaled ground-cuckoo 19–20"

Brazil

Tropical rain forest

Neomorphus radiolosus Banded ground-cuckoo 19–20"

Colombia, Ecuador

Lowland tropical rain forest, montane tropical rain forest

Neomorphus rufipennis Rufous-winged ground-cuckoo 19–20"

Venezuela, Brazil, Guyana, Colombia

Tropical rain forest

Neomorphus pucheranii Red-billed ground-cuckoo 18"

Brazil, Peru

Tropical rain forest

Sources: Haffner 1977; Phelps and de Schauensee 1978; Willis 1982; American Ornithologists’ Union 1983; Dunning 1987; S. N. G. Howell and Webb 1995.

Roadrunner Basics

9

southern New Mexico (Howard 1931), West Texas (Gehlbach and Holman 1974), and northeastern Mexico (Steadman et al. 1994). Recently, the westernmost specimen of Conkling’s roadrunner was found in Kartchner Caverns State Park in southeastern Arizona (Carpenter and Mead 2003). This roadrunner became extinct in the early Holocene (about six thousand years ago) when the climate was becoming warmer and drier. No recognized subspecies of the roadrunner exist today. Several ornithologists have examined the possibility of a modern, slightly smaller subspecies, with its distribution in the eastern part of the species’ range (Oberholser 1974; J. M. Hughes 1996a). Janice Hughes found that East Texas males were significantly smaller than California males in overall length and in wing, tarsus, tail, and bill lengths. Although such a subspecies is not recognized to date, perhaps a broadly clinal variation exists, with eastern birds being slightly smaller and darker and having larger white tips on the outer tail feathers than western birds (Pyle 1997).

Geared for the Ground Weighing slightly less than a pound, the roadrunner is nearly two feet in length, half of which is tail, and it stands nine to ten inches tall (plate 1). Adult weights range from ten to twenty-one ounces (J. M. Hughes 1996a). At a distance, the bird appears to have brownish, dull, streaked plumage, but at close range, its true colors show. The dark, white-fringed feathers are glossy and iridescent, with colors of olive, bronze, steel blue-black, brown or black, depending on the lighting. The upper breast feathers are a streaked mixture of buff, brown, black, and white, and the lower breast and abdomen feathers are pale gray. A large, white spot brightens the tips of all but the two center tail feathers (plates 4 and 8). There is also a white spot near the middle of each wing feather, which show when the wings are spread. These spots are visible on the wings of the sunning bird in plate 8. As expected for a ground-dwelling bird, a greater amount of the roadrunner’s body is devoted to moving on its feet than to using its wings. Almost a quarter of its body mass is in its leg muscles, and only about 14 percent in its wing muscles. The opposite is true for the average flying bird, which has about a quarter of its body mass devoted to flying muscles and 9 percent to leg muscles (Calder 1984). Comparing the body of the roadrunner with that of several of its tree-dwelling cuckoo relatives, ornithologist Andrew Berger (1952, 1954) found that the roadrunner’s pelvic bones were larger, its thigh muscles were better developed, and its breast muscles were less developed. He concluded that the major factors contributing to roadrunner speed are its very long legs and its alternate leg action. In addition, he found that several pelvic muscles originate farther forward and to the side in the roadrunner than in tree-dwelling cuckoos, improving the roadrunner’s balance during alternate leg movements. Figure 4 shows the leg and wing proportions of the roadrunner. The feet are about two and one-half inches long, with two toes pointing forward and two pointing backward, a pattern called zygodactyl (fig. 2). Unlike some birds that can move their outer toe either forward or backward as needed, the cuckoo’s outer rear toe is

10

The Real Roadrunner

Figure 4. Wing and leg of roadrunner. Adapted from Shufeldt 1886b.

permanently in the rear position. In tree-dwelling cuckoos, this foot pattern is considered an adaptation for perching. However, the muscle that controls the toe extension in the roadrunner is longer than that of tree-dwelling cuckoos, allowing the toes to spread out on a flat surface (Berger 1952). This trait doesn’t stop the roadrunner from using its feet for perching, and it spends considerable time in trees while foraging and nesting. The long cuckoo tail, useful for balancing on perches and flying, is also an asset to the roadrunner’s running movements. During nonstop straight running, the roadrunner holds its head and tail parallel to the ground, but it fans and tips its tail to maneuver during zigzag running and it lifts the tail to help brake. Other uses for the tail are as a foil when fighting, a brace and thrusting force when jumping, and a showy visual signal during courtship display. Several biologists, intrigued by this bird’s swiftness, measured its speed with a car and reported rates of fifteen to twenty miles per hour (Hunt 1920; Sheldon 1922; Cottam, Williams, and Sooter 1942). Researchers Kavanau and Ramos (1970) reported that roadrunners on an activity wheel could maintain running speeds of greater than eighteen miles per hour. These are extremely fast speeds for a bird lighter than a pound with feet that are only two and onehalf inches long. The much larger ostrich has been clocked at a running speed of thirty-one

Roadrunner Basics

11

miles per hour and up to forty-four miles per hour for short bursts. I marvel at the roadrunner’s ability to sustain its pace on the hot desert ground. The roadrunner can fly, but its flight is limited by its long, heavy legs and short, weak wings. It flies only short distances, usually starting from a fast run. Its typical flight is a wing-and-tail glide to the ground from a high perch, such as a cliff, a treetop, or a building. The longest flight I have witnessed was a glide from a high West Texas cliff across the Rio Grande onto the riverbank in Mexico, a ground distance of about a quarter mile. Roadrunners also glide short distances on the ground, taking off from a fast running start with wings furiously flapping. Unlike most other cuckoos, the roadrunner sports a large, moveable crest. This shaggy topknot is usually flattened (plate 1) but is raised when the bird is alerted, agitated, or courting (plate 2). The roadrunner’s most colorful feature is the inch-long patch of bare skin located around and behind the eye, extending to the back of the head (plates 2 and 3). This area is an indigo blue around the eye, and behind the eye it is white to bluish white with a blue edging. Following around the head, the color abruptly changes to a bright chrome orange. Unless the bird is agitated or courting, the orange area is usually not exposed, for it is covered by surrounding, small feathers (not by the crest, as has sometimes been described by other authors). A keen observer may detect a roadrunner’s level of agitation by paying attention to the amount of color patch that the bird exposes and the degree to which the crest is erected. The many possible crest positions can combine with differing amounts of color patch exposure to produce a variety of patterns. Some of these variations are shown in figure 5 and plates 2 and 3. These patterns could be signals in roadrunner language, but if so, their meanings are unknown to us. The color patch is most often exposed during the male’s courtship behavior, described in chapter 4. Another prominent head feature is the long, stout bill with a hooked tip, colored from dark charcoal to brown. It is a powerful one-and-a-half-inch weapon for capturing and killing prey (plate 6). The mouth of the adult is solid black, in contrast to the bright reddish pink mouth of the nestling. Roadrunner sexes are nearly impossible to tell apart in the field by appearance alone. Females average about an ounce lighter and are slightly under an inch shorter than males, but these differences are not discernible in the field. Folse and Arnold (1976) reported a subtle color difference in the bare skin around and behind the eye of roadrunners in South Texas. The females had a light blue to medium blue bare area behind the eye, whereas in the males the area was white or bluish white. This color difference was also difficult to discern, except at very close range and after having gained experience by observing a number of roadrunner pairs. Folse and Arnold cautioned that the color difference was based on only one population of roadrunners. They reported one successful nest where both parents had a white color behind the eye. Further study is needed on the amount of color variation overlap between the sexes and on the geographic variation of this feature. For both captive and wild birds, I was able to confirm the sex of birds only by their behavior during copulation in the breeding season. Individuals were identified by features such as missing tail feathers.

The Real Roadrunner

12

a

b

c

e

d

f

g

Figure 5. Variations of crest erection and color patch display: a, crest normal, color obscured; b, crest sleeked, exposed color; c, crest slightly erect, color exposed; d, crest moderately erect, color partially obscured; e, crest fully erect, color fully exposed; f, anterior view of e; g, posterior view of e.

At a distance, the roadrunner looks like the lesser roadrunner, but the former is two to four inches longer and has belly feathers that are grayish white rather than buff-colored, as in the lesser roadrunner. The lesser roadrunner’s foreneck and central chest area are not streaked, as they are in the roadrunner. If visible, the bare area behind the eye can help distinguish between the species, for it is blue, grading to crimson, in the lesser roadrunner.

Geographic Range The roadrunner is a nonmigratory species that normally moves only locally within a small area throughout its life. Despite this limitation, the species extended its range from its southwestern home into the Midwest and several southeastern states during the last century. Today it occurs in twelve states in the United States and in Mexico. The northern limits of its range extend through Northern California, southern Nevada, southeastern Utah, southeastern

Roadrunner Basics

13

Colorado, central Kansas, and south-central Missouri. (Records from the four-corners area where Arizona, New Mexico, Colorado, and Utah join are old and scant, and this region is not considered a part of the current range.) Eastern limits extend into central Missouri, eastern Arkansas, and northeastern, central, and southwestern Louisiana. The bird’s range also extends through much of northern and central Mexico, stretching from the Gulf of Mexico to the Gulf of California and across Baja California to the Pacific Ocean. The lesser roadrunner occurs in the northwestern part of the Yucatán Peninsula and the western areas of the countries of Mexico, Belize, Guatemala, El Salvador, Honduras, and Nicaragua. The roadrunner’s range overlaps slightly with that of the lesser roadrunner in a small slice of southwestern Mexico in the states of Sonora, Sinaloa, Jalisco, and Michoacán (S. N. G. Howell and Webb 1995). The roadrunner occurs at elevations ranging from 250 feet below sea level in Death Valley, California, to more than 11,000 feet high in the Rocky Mountains of Colorado. In most locations, however, it occurs below 7,000 feet.

Habitat Unlike most birds living in northern temperate climates, roadrunners do not migrate south in the winter but are permanent residents and breeders wherever they occur. In their yearround homes, they face the challenges of adapting to seasonal temperature changes and finding adequate food and shelter in the winter months. Thus they are limited to geographic areas that are favorable to both successful nesting and winter survival. That they have been able to expand their range and survive year-round in scorching deserts and areas with harsh winters says much about the adaptability of this bird. This habitat adaptability also suggests that the roadrunner cannot be characterized as an exclusively desert-dwelling bird, as it is popularly depicted. Major roadrunner habitat types include desert scrub, chaparral, savanna, open brushlands, open woodlands, and wooded stream corridors. They live in myriad plant communities, including paloverde desert scrub, blackbrush scrub, cholla grassland, chaparral, mesquite scrub, mesquite savanna, juniper savanna, piñon-juniper woodland, juniper-oak woodland, pine-oak woodland, oak-hickory woodland, cedar glades, streamside woods, and salt cedar thickets. Roadrunners even adapt to living alongside humans in less densely populated urban and suburban areas, such as yards, parks, agricultural lands, and vacant lots. They rarely occur in dense, unvegetated urban areas, dense brushlands, or woodlands with thick undergrowth. Considering the general vegetation structure of roadrunner habitats, all are similar—a mix of open area with aboveground, sturdy vegetation. The ground cover may range from bare ground to sparse, short bunchgrass or short lawn grass, while the taller vegetation may be cacti, small bushes, or trees. The open areas are essential for maneuvers to flush and catch fast-moving insects and other prey. Open areas are also important during nesting, for they provide nearby foraging

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The Real Roadrunner

places where the parents can monitor the eggs and the young. Tall vegetation or artificial structures provide a safe place for nesting and roosting.

Populations Roadrunners occur alone during the late fall and winter and in breeding pairs or small family groups during the breeding season. The roadrunner is not common anywhere the species occurs, but it is relatively more common in southwestern desert and brush habitats than in other parts of its range. The North American Breeding Bird Survey indicates that the U.S. roadrunner population as a whole has been stable or only slightly declining since the survey started in the 1960s, although local population fluctuations are common (Sauer, Hines, and Fallow 2003). Populations along the northern limits of the species’ range exhibit more fluctuations than elsewhere; fluctuations are associated with the severity of winter weather and the small numbers of individuals. The roadrunner is protected by the U.S. Migratory Bird Treaty Act of 1918, which gives legal protection to nearly all U.S. birds, but there are probably a few places where roadrunners are still killed for sport or out of the belief that they are a threat to quail and to domestic chickens (Stimson 1975; Meinzer 1993). The roadrunner is not federally or state-listed as threatened or endangered anywhere in its range. The Missouri Natural Heritage Database lists it as a species of conservation concern, with a rating of “sensitive 3” (rare and uncommon in the state, with between twenty-one and one hundred occurrences) (Missouri Department of Conservation 2003). The other northeastern and eastern states on the edge of the roadrunner’s range do not list the bird as a sensitive species (Arkansas National Heritage Commission 2002; Louisiana Department of Wildlife and Fisheries, n.d.; Kansas Bird Records Committee 2002). Two ground-cuckoo relatives—the banded ground-cuckoo of Colombia and Ecuador and the scaled groundcuckoo of Brazil—appear on the 2000 International Union for Conservation of Nature and Natural Resources Red List of Threatened Species (Hilton-Taylor 2000). One might ask why the roadrunner is surviving when many other bird species are in peril. Unlike specialist birds, the roadrunner is not tied to a specific habitat or food, giving it flexibility to adapt to changes that humans make to the environment. The cause of extinction for many animals is destruction of their habitat or food supply, but the roadrunner is omnivorous and can survive in many habitats, including urban areas. I think that other reasons for its success are its parental skills and its ability to nest as many as three times in a breeding season. Roadrunners are known to live for at least seven years in the wild. This longevity record is for a roadrunner that was banded as a nestling in central Oklahoma and found dead seven years and three months later only a few miles away from its hatching area (Patuxent Wildlife Research Center Bird Banding Laboratory, n.d.). A captive roadrunner lived to be nine years old in the Riverbanks Zoological Park in Columbia, South Carolina (Smith 1980). The U.S. longevity record for a cuckoo relative, the yellow-billed cuckoo, is five years in the wild.

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The discussion of the natural history of the roadrunner presented in the rest of this book includes these and many other facets of this creature that have contributed to its success. First, we will look at how the roadrunner meets the basic survival needs of finding food (chapter 2) and protection from the elements and predators (chapter 3). Chapters 4–7 describe the events during courtship and nesting that contribute to the reproductive success of this species. In chapter 8 we look at how the adaptable nature of this species has enabled it to expand eastward. Chapter 9 gives a glimpse of the rich folklore associated with this bird and the reasons it has been popular for centuries in the past as well as in modern culture.

2 Food and Foraging

atch the roadrunner for an hour at his daily business of catching food and you will deem him among the most amazing of all the desert’s amazing creatures.” Thus did Oklahoma ornithologist and bird artist George Miksch Sutton (1940, 37) pay homage to one of his favorite birds, zeroing in on what has fascinated many of us about this bird. Indeed, we must admire a bird that casually eats scorpions, tarantulas, bees, wasps, ants, centipedes, rattlesnakes, and horned lizards. Much of the folklore about the roadrunner is related to its eating venomous creatures (chapter 9). It has also been lauded for eating numerous crop-damaging insects, such as grasshoppers and caterpillars (Anthony 1897; McAtee 1931). But the roadrunner has not always been appreciated for its diet. In the early 1900s several states paid bounties for its destruction because it was accused of eating young chickens and quail (McAtee 1931). Hunters blamed it in years when quail populations were low. Texas folklorist J. Frank Dobie lamented this persecution, pointing out that roadrunners ate snakes and rats, both greater threats to quail populations than are roadrunners. Observing abundant roadrunner and quail populations in the same areas, he commented: “You will not find the most shotgun hunters and the most quail in the same pasture” (Dobie 1939, 8). On the other hand, several biologists, including well-known naturalist Aldo Leopold, observed roadrunners killing baby quail (Leopold 1922; Law 1923; R. E. Wright 1973). Several government studies helped dispel the myth of the roadrunner as a major quail predator, showing instead the benefits of its insect eating, and the bounties on roadrunners were lifted. The U.S. Department of Agriculture’s former Bureau of Biological Survey

W

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The Real Roadrunner

conducted a study of roadrunner stomach contents from 1885 to the early 1900s. Collected from seven southwestern states, these unpublished data provided an unbiased look at the roadrunner diet (Patuxent Wildlife Research Center, n.d.). Similar stomach contents studies in Southern California by Harold Bryant (1916) and in Arizona by D. M. Gorsuch (1932) also addressed the roadrunner’s reputation as a killer of chickens and quail. Together these studies examined the stomach contents of more than two hundred roadrunners. Information from these studies, as well as from several published accounts and my unpublished observations of the roadrunner diet, serve as the basis for the following section on the diet of the adult roadrunner.

Adult Diet The roadrunner may be characterized as an opportunistic omnivore—that is, it eats any edible item it encounters that it can catch, kill, and swallow. Such items include small animals of all kinds, eggs, fruit, seeds, and carrion. Thus, the roadrunner’s diet is a good reflection of the food available in its habitat throughout the year. This adaptable diet has been a factor in this nonmigratory bird’s winter survival, expansion into new habitats, and success in urban areas. Insects and other arthropods are the year-long diet staples of the adult roadrunner (about 80 percent of the food volume). The most common insect foods are grasshoppers, beetles, and crickets, but it also eats caterpillars, bugs, ants, bees, wasps, fly larvae, butterflies, and moths. Other invertebrate foods are scorpions, whip scorpions, spiders (including tarantulas), sow bugs, centipedes, millipedes, and land snails. Snails in the roadrunner diet are especially well documented (Baird 1859; Wheelock 1904; F. M. Bailey 1917; Dobie 1939; Cutright and Brodhead 1972; R. E. Wright 1973; Kazmaier, Ruthven, and Synatzske 1999; author’s observations). Although insects and other arthropods are not very active in the winter, the roadrunner somehow manages to find them. Items found in the stomachs of winter roadrunners have included spider eggs, European mantid (praying mantis), pupal stages of moths and butterflies, and remains of insect egg cases. Vertebrates accounted for only a small part of the food volume in stomachs (about 16 percent), but almost half of the roadrunners had eaten vertebrates. Fewer reptiles, birds, and amphibians appeared in stomachs in winter months than in summer months, but the roadrunners ate about the same amount of small mammals throughout the year. Roadrunners eat more lizards than any other type of vertebrate. Lizard prey include fence, side-blotched, spiny, alligator, collared, horned, and prairie lizards, whiptails, and skinks. The only turtle reported from a roadrunner stomach was a yellow mud turtle hatchling (Kazmaier, Ruthven, and Synatzske 1999). James Cornett (1999) observed a roadrunner attack on a juvenile desert tortoise, but the bird gave up after the tortoise withdrew into its shell. Despite the roadrunner’s common name of “snakebird” and reputation for eating snakes, snakes are only a small fraction of the bird’s diet. Roadrunners have been observed killing

Food and Foraging

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venomous snakes, such as the western diamondback rattlesnake and the western coral snake. Other reports of roadrunner stomach contents have included racer, blind, garter, patchnose, whip, rat, and ground snakes. An unusual find was a thirty-inch-long rattlesnake in an Arizona roadrunner’s stomach (Gorsuch 1932). However, the snake could have been dead when the bird found it, for roadrunners commonly eat dead prey. To determine how large a snake roadrunners would attack, Wyman Meinzer (1993) presented live rattlesnakes to wild roadrunners in northern Texas. These roadrunners would not attack a rattlesnake more than two feet long, retreating whenever a snake became aggressive. However, I have seen roadrunner parents defend their nest by attacking and chasing away rattlesnakes and other snakes that are longer than two feet (see chapter 5). Although only a few amphibians are reported from roadrunner stomachs, the birds probably eat toads and frogs in habitats where these are common. A study in north-central Texas reported frogs and toads in four of the sixteen roadrunner stomachs that were examined (Parmley 1982). The roadrunner eats a variety of bird species. Adult birds captured live by roadrunners include black-chinned hummingbird, swift, orchard oriole, house finch, Oregon junco, and house sparrow (W. E. Allen 1932; Sutton 1940; Zimmerman 1970; Binford 1971; Spofford 1976). A northern cardinal, a dark-eyed junco, and a pine siskin caught in a mist net fell prey to a roadrunner (Barclay 1977). Roadrunners have been reported to capture young quail and to rob nestlings from nests of mockingbirds, house finches, Abert’s towhees, and purple martins (Bendire 1895; W. K. Fisher 1904; J. M. Hughes 1966a; Finch 1981). Bird species found in roadrunner stomachs include northern bobwhite, Inca dove, Bewick’s wren, cactus wren, meadowlark nestling, canyon towhee, pyrrhuloxia, lark bunting, Oregon junco, chipping sparrow, unidentified warblers, and unidentified bird eggs (Bryant 1916; Gorsuch 1932; Kazmaier, Ruthven, and Synatzske 1999; Patuxent Wildlife Research Center, n.d.). The mammals that roadrunners eat comprise both diurnal and nocturnal animals, including bats, rabbits, ground squirrels, antelope squirrels, gophers, kangaroo rats, cotton rats, wood rats, pocket mice, harvest mice, white-footed mice, and voles. Plant food was only a small part of roadrunner stomach contents. However, plant food appeared to be an important supplement in the fall and winter in some areas, constituting from 6 to 10 percent of the food volume. About twice as much plant material occurred in stomachs in the winter as in other seasons, especially sumac berries and prickly pear cactus fruits. Other seeds or fruits were from the following plant groups: grass, legume, nightshade, amaranth, aster, geranium, goosefoot, mallow, milkweed, morning glory, poppy, purslane, rose, smartweed, and spurge. It is possible that roadrunners did not directly eat all of these plant materials, for some might have been in a prey animal’s digestive tract. However, fruits and seeds were in larger quantities than could be accounted for in this way alone. One roadrunner stomach was filled with nothing but oat seeds and plant tissue. Several observers, including the author, have watched wild roadrunners eating sumac berries and cactus fruit.

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The Real Roadrunner

Roadrunners are notorious for eating carrion, pet food, garbage, and just about anything edible they come across. One nesting female I followed in West Texas ate at least twenty puffed cheese snacks she encountered. Several pairs I watched in urban areas depended on pet food for themselves and their nestlings.

Nestling and Juvenile Diet Even though no stomach contents studies of nestlings have been reported, field observations provide a good picture of nestling diets. The information in this section comes from studies at four nests in Arizona (Ohmart 1973); at eleven nests in Oklahoma, New Mexico, and Texas by the author; and at several nests in northern Texas (Meinzer 1993). At the first nest I watched, the difference between what the parents ate and what they fed their nestlings was striking. As the parents foraged, they snapped up insects along the way, but after catching a lizard or a bird, they ran back to feed it to a nestling. As I watched additional nests, I realized that vertebrates were the major nestling food. Feeding a vertebrate to a nestling makes sense, for it provides more food per trip than would a small insect. Ohmart (1973) reported that lizards were the main nestling food at four Arizona nests. The ability of a parent to capture large numbers of vertebrates to feed nestlings, while it continues eating insects, must be important to successful roadrunner nesting. At my eleven nests, vertebrates made up the majority of the food, although insects were also a staple. I estimated that about 90 percent of the food volume brought to the nest was vertebrate. Lizards were fed at all nests, snakes and rodents at five nests, and birds and amphibians at four nests. One of the snakes was a fifteen-inch-long rattlesnake fed to a nestling in New Mexico. A roadrunner parent fed a small massasauga rattlesnake to its fledglings in northern Texas (Meinzer 1993). Despite these observations, we cannot accept the generalization that reptiles are the main food at every roadrunner nest. The opportunistic nature of the roadrunner allows it to take advantage of whatever food is abundant during nesting. For instance, parents at the two urban nests I watched fed more birds and insects and fewer reptiles to their nestlings than did parents at rural nests. House sparrows, bird eggs, pet food, and human-provided meat strips were important food items at these nests. Wyman Meinzer (1993) reported several instances of opportunistic nestling feeding at nests in northern Texas. At one nest the Texas horned lizard was the major nestling food, in an area where these reptiles were abundant. Later in the breeding season at a nest in the same general area, most nestling food was crickets and grasshoppers, coinciding with the height of a grasshopper infestation. Even when Meinzer brought a horned lizard to the nest site, the parents ignored it, in the midst of the plentiful insect supply. At another nest in the area late in the breeding season, almost all of the food items brought to the nestlings were insects. All these feeding observations are from Southwestern nests, where reptiles are abundant during most of the breeding season. Unfortunately, no nestling food data exist from northern

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or eastern parts of the roadrunner’s range. An important contribution to the natural history of the roadrunner would be observations on nestling diets in Kansas, Missouri, Arkansas, Louisiana, Colorado, Utah, and Northern California.

Quantities of Food I am amazed at how much a roadrunner can cram into its stomach. For instance, Harold Bryant (1916) reported that a male roadrunner from Southern California had the following items in his stomach: 1 horned lizard 4 small lizards 12 small grasshoppers 4 large grasshoppers 50 tenebrionid beetles l weevil 10 carabid beetles 1 beetle larvae 2 moths The quantity of venomous animals that one roadrunner can eat at a sitting is remarkable. One stomach contained 250 wasps, and another had 7 scorpions (Patuxent Wildlife Research Center, n.d.). No evidence has been found that roadrunners suffer ill effects from eating venomous animals.

Foraging Areas Correlated with its diverse diet, the foraging areas of the roadrunner are myriad, including open ground, underground, midair, pavement, grass, shrubs, trees, water, and human structures. Animal trails or other open pathways are major roadrunner feeding areas, for these provide room for it to freely walk, run, and maneuver to flush and catch prey. Besides eating insects and other small animals along a trail, roadrunners also pick up insects from the animal dung found there. Paved roadways are also major foraging areas, probably accounting for the demise of a few roadrunners. Ornithologist James Cornett (2000) observed roadrunners feeding on live grasshoppers on pavement, where these insects did not appear to be as camouflaged as they did on the nearby desert soil. I have often seen the birds feeding on squashed insects and other carrion on pavement. Foraging roadrunners frequently dig in the ground for food. Desert naturalist Edmund Jaeger (1947) encountered one turning over large plates of caked, dry mud with its bill, looking underneath for insects. In less than ten minutes, this bird had turned over more

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The Real Roadrunner

than thirty of the seven-inch-wide mud plates, finding many insects beneath them. I have watched roadrunners dig with their bills in sandy soil or loose dirt, throwing sand afar, especially when digging in the funneled holes of ant lions. Sometimes roadrunners forage in trees or bushes rather than on the ground. Amid the foliage and branches they find such delicacies as caterpillars, daddy longlegs, European mantids, bird eggs, and bird nestlings. At one West Texas nest I watched, arboreal feeding was especially common at times when the nearby cottonwood trees were loaded with caterpillars.

Foraging Methods Because of the many types of food that the roadrunner eats, the bird needs to be a jack-ofall-trades when it comes to techniques for detecting and capturing prey. Its most common foraging mode is a slow walk, interspersed with brief stops while it peers into, under, and on top of everything along its pathway. When it spots prey, the capture happens so quickly that observers get only a partial glimpse of the technique and skill involved. During my field studies, I was often able to follow closely behind foraging roadrunners, for they quickly adjusted to my presence. Even when I was as close as ten feet, however, I never saw the prey before it ended up in the bird’s bill. One clever way that the roadrunner forces prey from its hiding place is to startle it by wing-flashing. Slowly walking or standing still, a roadrunner will quickly thrust its wings outward and then back to its body. As the wings extend outward, the wing feathers are almost fully spread, revealing the white patches near the center of each feather (fig. 6). The sudden movement and the flash of white cause insects and lizards to scurry from hiding places, only to be snapped up by the waiting bird. I have watched wild roadrunners do this in many different habitats. Juvenile roadrunners as young as thirty-five days old wing-flash as they begin to forage for food on their own. My hand-reared birds would flash their wings as they hunted in the short grass in their large outdoor enclosure. In tall, thick grass, the roadrunner has little room for ground maneuvering or wingflashing. Here the bird shifts to another foraging mode. Wyman Meinzer (1993) watched a wing-flapping, tail-fanning, leaping technique used by a roadrunner to flush insects in tall grass. Another flushing method the bird uses in tall grass is to brush the grass with its head held low to the ground, forcing insects to move. Other birds that wing-flash to flush insects—such as the pheasant cuckoo, the northern mockingbird, and the American redstart—also have white patches on their wings (Selander and Hunter 1960; Sieving 1990). However, some birds that wing-flash do not have wing patches, including some tropical American mockingbirds, the gray catbird, and the brown thrasher. The roadrunner takes every opportunity to let others do the work of flushing or capturing prey for it. Several observers have reported this “lazy” way of getting food. Ornithologists Florence Merriam Bailey (1922, 1923) and Sally Spofford (1978) reported roadrunner attacks on live birds caught in traps. One naturalist endured a roadrunner stealing small

Food and Foraging

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Figure 6. Roadrunner flashing its wings to flush prey.

birds he had shot before he could retrieve them himself (A. B. Howell 1916). A bird bander using mist nets to catch birds unintentionally provided several meals for a roadrunner (Barclay 1977). In South Texas a roadrunner was seen moving along with a white-tailed deer herd that was feeding in low vegetation (Michael 1967). Closer observation revealed that the roadrunner was grabbing insects stirred up by the deer. This behavior is similar to that of cattle egrets that associate with cows and other large, grazing animals to eat insects found on and around them. The scissor-tailed flycatcher follows wild turkeys and feeds on insects flushed by them. Roadrunners will even follow humans to find prey. In the mesquite grasslands of South Texas, a common ranching practice is to burn the needles from prickly pear cacti to allow cattle to eat the juicy pads and fruity without getting thorns. One rancher reported that a roadrunner followed the pear-burning worker every day, searching the singed cacti for roasted worms and bugs (Dobie 1939). In farming country, roadrunners will follow a plow to grab insects and worms exposed in the newly turned soil (Dobie 1939). The roadrunner may wait, catlike, in ambush outside animal holes or near bird feeders until unsuspecting prey appears. A black-chinned hummingbird met its end when a roadrunner caught it in midair at a hummingbird feeder that ornithologist Sally Spofford (1976) was watching. Ornithologist Dale Zimmerman (1970) reported a roadrunner lying in wait in a juniper for house sparrows in an area where they usually congregated. Another observer reported roadrunners catching a house finch and two young California quail by lying in wait at a bird feeder (R. E. Wright 1973). There are many popular accounts of bird-watchers supplying meat or dog food to their resident roadrunner to the extent that the bird sometimes becomes dependent on this food

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The Real Roadrunner

source. I observed this in New Mexico, where a homemaker daily put out strips of raw chicken for a roadrunner pair that were nesting in a yucca in her yard. The birds would grab the meat as soon as she moved away a few feet. When their eggs hatched, they took the chicken, along with food from the dog dish, to feed the nestlings. Even in areas far from human influence, roadrunners quickly learn to panhandle. One regularly came to a camp in a New Mexico wilderness area to eat the entrails of small animals that biologists were skinning (F. M. Bailey 1922). I was amused to hear from an Arizona man whose wife made small meatballs for their resident roadrunner to eat. After the bird began nesting nearby, it took these meatballs, often two at a time, to feed the nestlings. The meatballs disappeared so fast that his wife started freezing them in large quantities to keep ahead of the roadrunner’s demands. Soon the bird brought its newly fledged juveniles for the treats. Many people have written me about wild roadrunners regularly coming by their home for a handout.

Killing and Swallowing Large Prey Unlike birds of prey that use their feet to kill, tear, or handle their prey, roadrunners stun and crush prey only with their bill, and they swallow it whole rather than in pieces. Biologists who have examined roadrunner-killed animals report blows to the base of the skull as the cause of death (Anthony 1896; Bleich 1975). Landing a snake for dinner, especially a poisonous one, is one of the greatest tests of the fighting skill and mettle of the roadrunner. The bird first slowly circles the snake sideways with tail fanned and tipped toward the snake, then jabs at the snake’s head with lightning speed. The bird will often flash its wings rapidly, revealing white patches, as it circles the snake. The northern mockingbird also flashes its wings, showing a white patch, in the presence of snakes (Selander and Hunter 1960). When the snake strikes back, the faster-moving roadrunner leaps out of reach of the strike, then begins jabbing at the head again. Another maneuver of the roadrunner is to hold its wings fanned outward, like a bullfighter’s cape, to divert the snake’s strike. The final kill is made by repeated jabs to the snake’s head. Sometimes a pair will work together in a team attack, with both circling the snake until one gets an opportunity to jab at the snake’s head. Before swallowing large prey, the roadrunner usually beats it repeatedly on a rock or other hard surface. Lizards, snakes, birds, and small mammals may be beaten for as long as thirty minutes. The research of Beal and Gillam (1979) found that this beating behavior resulted in broken bones and an elongated body, making the prey easier to swallow than an intact body would be. My hand-reared birds would kill and beat the live vertebrate food I gave them. They would even vigorously beat a two-inch-long strip of meat before they swallowed it. Bird prey receives the additional treatment of feather plucking. Most of the bird’s wing and tail feathers are pulled out as the roadrunner holds the end of each feather in its bill and rapidly shakes and beats the bird’s body.

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After all the beating and feather plucking, the roadrunner is ready to swallow the prey headfirst with several body-heaving gulps. Swallowing may take more than five minutes for large lizards, rodents, and birds. Even after swallowing, there may be a lump in the bird’s throat for a long time, as digestion and more swallowing move the food along. Several sightings of a snake hanging out of a roadrunner’s bill for an hour or more have been reported. While swallowing very large prey, the roadrunner is temporarily immobilized and possibly vulnerable to predators. Ironically, parents don’t spend as much time preparing the food for their nestlings as they do for themselves. Parents rarely beat or pluck feathers from nestling food. I have seen roadrunner parents taking unplucked house sparrows to the nest, even though the adults plucked the feathers of the sparrows they ate themselves.

Special Case: Swallowing Horned Lizards Of all the food a roadrunner eats, the horned lizard merits special attention because the large, hard horns on its head and body pose a peril during swallowing. Herpetologist Wade Sherbrooke raised several roadrunners to study their method of killing and swallowing horned lizards. He was kind enough to discuss his research with me and provided plates 5 and 6, which show a roadrunner swallowing a Texas horned lizard. Sherbrooke observed that roadrunners would attack and eat the smaller, less spiny round-tailed horned lizard more often than they would attack the larger, spinier Texas horned lizard. Also, the Texas horned lizard would inflate its body when attacked by a roadrunner, making its horns stand out more than normal. Wyman Meinzer (1993) also reported roadrunners backing off from large Texas horned lizards, notably on one occasion when the horned lizard squirted blood from around its eye area. Sherbrooke described the roadrunner’s technique for beating a horned lizard: “The bird grasps any leg or the tail in its beak, rapidly raises its head and neck (tilting its head to one side) and swings the outwardly forced body of the prey upwards and around through a vertically extended, oval-shaped trajectory that culminates in a downward accelerating slam against the ground” (1990, 172). One beating that he watched lasted more than five minutes, with about twenty-one slams per minute, although it didn’t break the lizard’s horns. Several times he saw a half-swallowed Texas horned lizard being regurgitated and beaten again before being finally swallowed. The horned lizards were swallowed headfirst and upside down. In this position the horns face outward rather than toward the roadrunner’s internal organs. Even when a large horned lizard is upside down, swallowing it is a hazardous undertaking for a roadrunner. Recently, a dead juvenile roadrunner was discovered in Texas with a large Texas horned lizard half-swallowed in its throat. Even though it had swallowed the lizard upside down, the juvenile’s neck was split open on one side, apparently by one of the lizard’s horns (Holte and Houck 2000).

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The Real Roadrunner

Treatment of Small Prey Roadrunners attack scorpions and wasps at the tail end, whereas tarantulas are attacked at the head (Sutton 1913). Roadrunners flick and eat the legs from tarantulas and pull the pinchers from crayfish before eating them. On the other hand, the birds gobble up many types of stinging animals without any prelude, including bees, wasps, and scorpions. But thorny plant food takes a little preparation. One roadrunner rolled around a tasajillo cactus fruit on the ground before eating it, presumably to knock off the spines (Meinzer 1993). Roadrunners beat snail shells on a rock or other hard surfaces before they pluck out and swallow the soft body. In West Texas, I saw the same rock repeatedly used by roadrunners for beating snails, with a heap of shells around it. Robert Wright (1973) observed roadrunners in California hammering snails against rocks and eating the soft bodies or feeding them to nestlings. At times, roadrunners may also eat the shells as well as the soft body parts, as do some other birds. The shells may be used as a source of calcium by female birds during egg laying.

Food Pellets Vertebrate-eating birds such as eagles, hawks, owls, and vultures are well known for regurgitating pellets containing undigested hair, bone, teeth, nails, and other animal parts. Many invertebrate-eating birds also produce pellets containing insect exoskeletons, mollusk shells, and grit. By identifying the animal parts in these pellets, ornithologists get clues to the bird’s diet. Adult roadrunners have never been observed producing pellets in the wild, although I don’t doubt that they do, especially when their diet is high in vertebrates. The nestlings I raised on a vertebrate diet did not produce pellets until the birds were about one month old, when they regularly regurgitated pellets containing undigested bone fragments, toenails, hair, and insect exoskeletons. A typical pellet was about three-fourths inch long by one-half inch wide. These birds continued to regurgitate pellets as adults, the quantity depending on how much vertebrate food they ate. The ability of the nestling roadrunner to eat and digest vertebrates without producing pellets would be an interesting area of study.

Food Habits of Ground-Cuckoo Relatives Little has been recorded about the food habits of the lesser roadrunner and other groundcuckoos. The diets of ground-cuckoos include large insects, scorpions, centipedes, spiders, small frogs, birds, lizards, and occasionally fruit—a diverse diet reminiscent of that of the roadrunner. Stomach content data exist for one lesser roadrunner (stomach filled with grasshoppers), two rufous-vented ground-cuckoos (scorpions, roaches, unidentified insects, and seeds), and several pheasant cuckoos (grasshoppers, cicadas, small lizards, and nestling birds) (A. H. Miller 1932; Wetmore 1968; Stiles, Skutch, and Gardner 1989).

Food and Foraging

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A ground-cuckoo relative, the pheasant cuckoo, flashes its wings when foraging for insects, revealing small, white wing patches (Sieving 1990). The striped cuckoo flushes prey by flashing the feathers of its alulae (the processes of bird wings that correspond to the thumb) like fans, as it sways from side to side (Stiles, Skutch, and Gardner 1989). Like the roadrunner, ground-cuckoos are adept at using other animals to flush food. The lesser, red-billed, and rufous-vented ground-cuckoos follow large, fast-moving army ant swarms to feed on the flushed insects and other small animals (Willis 1982; Greene, Wilcove, and McFarland 1984). The rufous-winged ground-cuckoo is called pàjaro vaquero, or cowboy bird, because it often follows bands of peccaries (piglike, hoofed mammals), as a cowboy might follow a herd of cattle (Haffer 1977). Various other local names for groundcuckoos throughout their range mean “peccary bird.” In tropical forests, both red-billed and rufous-vented ground-cuckoos follow on the ground below monkey troops moving through the trees. The birds feed on the flushed insects, dropped fruit, and fruit remains in the monkey droppings (Siegel, Hamilton, and Castro 1989).

3 Daily Survival

he previous chapter showed the opportunistic nature of the roadrunner’s feeding habits, which allows it to survive in many habitats throughout the year. Now we will look at other adaptations of this bird that help it survive in the extremes of hot southwestern summers and cold midwestern winters.

T

Surviving Hot Summer Days Reducing Activity Watching a roadrunner scurrying about on the scorching desert ground, we marvel at its survival in temperatures that quickly stress humans. Yet the roadrunner thrives in the hottest areas in the United States, not only in the hot, dry desert summers but also in the hot, humid midwestern summers. Most other ground-dwelling desert animals, such as rodents, rabbits, and scorpions, take the easier route of being active during the cooler night or twilight hours and resting in the comfort of their cool, humid burrows during the daytime. Desert-dwelling diurnal birds survive by avoiding activity during the hottest times of day, staying off the hot ground, and sometimes flying long distances to find water. Some desert birds also have nasal salt glands that help them conserve water. Because the roadrunner is a diurnal, ground-dwelling bird that survives extreme desert conditions, its temperature control and water balance have received attention from avian physiologists studying desert survival (W. R. Dawson and Schmidt-Nielsen 1964; Calder 1966, 1967b; Calder and Schmidt-Nielsen 1967; Lowe and Hinds 1969; Ohmart 1972).

30

The Real Roadrunner

Their research results suggest that, like other desert birds, the roadrunner primarily survives desert heat by avoiding activity during the hottest times of day and by conserving water via its nasal salt glands. Though it may forage off and on throughout the day, a roadrunner reduces its activity and takes rest stops during the peak hot period from noon to late afternoon. While standing in a shady area, it rapidly pants and flutters its throat skin (Calder 1967b). It also holds its wings down and out, allowing body heat from its sparsely feathered sides to escape into the air. Wild roadrunners I watched in Texas, New Mexico, and Oklahoma all had this activity pattern on hot summer days. The exceptions were parents that were feeding nestlings. To meet the demands of their hungry nestlings, parents often foraged through the hottest times of the day. Since birds do not have sweat glands, they cool off by water evaporation from respiratory surfaces and from their skin. Roadrunners and some other birds also increase the evaporation rate by fluttering their gular areas, the thin-skinned floor of their mouth and upper throat. Blood vessels in this area become engorged during high temperatures, losing heat into the air. This works to control body temperature, except in extremely hot air temperatures (about 120°F and higher). Another way birds lose heat is by an increased blood flow to the legs and feet, again releasing heat from the body. The arteries and veins in the roadrunner’s neck and head are arranged to help keep the brain from overheating (Kilgore, Bernstein, and Hudson 1976). The carotid network of veins and arteries is an effective site for heat exchange between warm arterial blood flowing toward the brain and cool venous blood returning from the face and the nasal and throat areas. Birds also are able to survive hot days because they tolerate body temperatures up to 7°F above their normal body temperatures (W. R. Dawson and Schmidt-Nielsen 1964). In addition, the normal body temperature of many birds, including the roadrunner, is higher than that of humans, giving them greater leeway in hot weather. The roadrunner’s normal body temperature is 101–104°F. What does the roadrunner do when no shade is available? By turning its back to the sun, it positions its head, neck, outspread wings and tail, and body to shade its gular area and feet. I often observed wild and captive birds standing this way in the hot sun. The roadrunner also stands this way at the beginning of a light rain, allowing water to cover the tail and wing feathers. My captive birds also assumed this posture when lightly sprayed with a water hose on hot days. Another way in which the roadrunner avoids heat is by getting off the ground, which may be many degrees warmer than the surrounding air. Wild roadrunners often spend the hottest time of the day perched in a tree or bush above the hot ground. A negative side effect of the panting/gular flutter method of body cooling is that it requires a lot of water. So ultimately, heat survival depends on offsetting this water loss to maintain the right salt-to-water balance in body fluids. The roadrunner is well equipped for this task, as described in the next section.

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Conserving Water Drinking large amounts of water is the simplest way to restore water balance, and the roadrunner drinks water whenever it is available in the bird’s habitat. Also, the roadrunner’s animal diet, which is from one-half to two-thirds water by weight, can provide most of the water that the bird needs (Calder 1968). The roadrunner conserves water through its nasal salt glands, which remove sodium and potassium salts from its bloodstream and eliminate these salts in droplets from its nostrils (Ohmart 1972). This avoids the water loss that would occur if salts were excreted in its urine. Although all birds have salt glands, these glands are better developed and functional in birds living in marine and desert habitats. Those marine and land birds that have active salt glands, including the roadrunner, also have more kidney tissue for their size than do other birds (M. R. Hughes 1970). Nestling salt glands, which are relatively larger than those of the adult, are discussed in chapter 7. In low humidity, nasal droplets quickly dry and leave white, crusty salt deposits in a roadrunner’s nostrils and on the bill. In high humidity, however, the salt crusts are not noticeable, for the drops fall off the bill before they have time to dry. Adults that I kept outdoors in hot, humid Oklahoma summers never had white salt deposits around their nostrils. Concentrating urine before it leaves the body is a way that some desert mammals conserve water. Although initial research showed that the kidneys of water-deprived roadrunners could not concentrate urine any better than those of nondesert birds (Calder and Bentley 1967), it was later found that after urine leaves the roadrunner’s kidneys, some water may be reabsorbed from it during hot, dry times (Ohmart, McFarland, and Morgan 1970). During the nesting season, the parents sometimes eat their nestlings’ fecal sacs, which have a high water content (chapter 6).

Surviving Cold Nights Given all of the roadrunner’s ways to avoid heat stress, it seems paradoxical that the bird also needs ways to keep warm. Cool temperatures pose a problem for the roadrunner not only in midwestern winters but throughout the year wherever it occurs. People who live in the desert know that even on hot summer days the temperature in the dry air falls quickly after the sun sets, often falling as much as 40°F below the daytime high. How do cold temperatures affect the roadrunner? The energy from food is needed for fuel to keep the bird’s temperature steady when the air temperature is below its normal body temperature of 101–104°F. So the roadrunner simply goes into a hypothermic state— that is, it drops its body temperature—to conserve food energy at night. This process is similar to turning down a thermostat on winter nights to save fuel costs. The body temperature of a roadrunner may drop 7 to 15°F at night as the surrounding temperature falls (Ohmart and Lasiewski 1971; Vehrencamp 1982). This energy savings comes with a cost to the lives of some roadrunners, for having a low nighttime temperature means that the bird is sluggish and slow to react to predators.

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The Real Roadrunner

After my captive roadrunners had been roosting for several hours, I could enter their outdoor cage and take them off their perch, with no visible response from them. James Cornett (2001) reported similar nighttime lethargy for the roadrunner he raised. Soon after leaving the roost, a roadrunner basks in the sun to warm up quickly. With its back to the sun, it raises its scapular feathers, exposing a patch of dark, bare skin that absorbs the sun’s heat, much like the operation of a solar panel (plate 7). This sunning behavior can raise the roadrunner’s body temperature about 5°F and saves the bird about 550 calories per hour (Ohmart and Lasiewski 1971). A roadrunner will sun in this position periodically throughout the day during cold weather. The roadrunner also suns itself by lying flat on hot ground or pavement at various times throughout the day. From a standing or walking position, the bird suddenly drops prone to the ground, spreads its wing and tail feathers, and exposes the bare, colored skin behind the eye (plate 8). It stays in this posture for several minutes and appears to be in a daze. I could approach wild roadrunners in this sunning posture without their seeming to notice me. Although I saw wild roadrunners sun this way on warm, sunny days, my captive birds sunned this way on cold, sunny days as well. Several ground-feeding songbirds, including northern cardinals, blue jays, towhees, and thrashers, sunbathe in a similar prone position on hot surfaces. Whether or not this behavior is related to the control of body temperature is not known. Several biologists have suggested that this behavior may be for getting rid of feather mites and other parasites. While sunbathing, the roadrunner, like other birds, may dust-bathe and preen its plumage. Sand bathing may also help dislodge flea, mite, and lice parasites. The ability to lower their body temperature at night is beneficial to roadrunners that live in cold climates, where the birds have to contend not only with low temperatures but also with long periods of food shortage, as discussed in the following section.

Surviving Winter Food Shortages Wherever the roadrunner occurs, a crucial factor in the bird’s success is its ability to find winter food, given that this bird does not migrate. Insects and reptiles are usually dormant during winter, even in the Southwest. Some will “come to life” and move around sluggishly on warm days in winter—easy marks for a hungry roadrunner. However, these warm days are rare and would not provide enough steady food throughout the winter, even in southwestern habitats. In the same manner that the roadrunner drops its body temperature each night, it may also become hypothermic to save energy on very cold, dark days in periods of food scarcity. George Miksch Sutton (1986) suggested this possibility after seeing few roadrunners in winter during his many years of bird study in Oklahoma. James Cornett (1998, 2001) proposed this as one reason why roadrunners are scarce in southwestern areas during cold periods. Whether the roadrunner lowers its metabolism to the point of torpor, in which a bird’s thermoregulatory system shuts down, is another question. The lack of roadrunner sightings

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in winter could be explained by the bird’s becoming inactive and temporarily hypothermic rather than becoming torpid when food is scarce. The poorwill, a nonmigratory, insect-eating bird, becomes torpid during winter when food is scarce. Desert biologist Edmund Jaeger (1948, 1949) monitored a wild poorwill that stayed in such a torpor from mid-December until late February, a period during which there were no flying insects. Later study of captive poorwills found that less than one-half ounce of stored fat could sustain a poorwill in a torpid state for one hundred days, if the temperature stayed at least as low as 50°F. This state was not induced by cold, but rather after the birds lost 20 percent of their weight. Another bird that feeds on insects as it flies, the white-throated swift, also enters a torpid state during times of low food supply. No one has observed the roadrunner in a torpid state such as that reported for the poorwill. Unlike the poorwill or the swift, the omnivorous roadrunner is not limited by the type of food available in an area. Nor is it limited to eating only live food. Stomach contents and observations of roadrunners in Oklahoma winters suggest that the birds eat insects, arthropods, insect egg cases, seeds, fruit, small songbirds, and carrion during severe winter weather and probably eat small rodents as well (Geluso 1969, 1970a, 1970b; Shetlar 1971; Sutton 1973). But finding this food in areas where it is scarce or in long periods of extreme cold or snow cover is another matter. Snow cover not only buries food but also makes it difficult for a ground-dwelling bird to move about and forage for food. An interesting question is where the roadrunner forages in the winter, especially in areas with prolonged low temperatures and snow cover in the northern parts of their range. Some clues come from bird-watchers in these areas. A northwestern Arkansas bird-watcher informed me of seeing a roadrunner kill European starlings in his yard during extremely cold winter weather with prolonged snow cover. This was the first time he had seen a roadrunner in the area, and it disappeared after the weather improved. Another roadrunner enthusiast from Arkansas wrote me about a lone roadrunner that came into his yard only during long periods of severe winter weather, readily taking food from him. Feeding by humans during periods of severe winter weather may keep some roadrunners from starving. In the Oklahoma panhandle, George Miksch Sutton (1967) observed roadrunners moving to bases of cliffs in severe winter weather for shelter and possibly for insect food among the rocks strewn at the base of the cliff. Kathleen Beal (1981) observed roadrunners foraging in tall grass during severe winter weather in south-central Oklahoma. Another way the roadrunner may survive periods of food scarcity is by living off its stored body fat. Kenneth Geluso (1969, 1970a) reported that the eight Oklahoma roadrunner specimens he examined in early winter were heavier and had more lump fat than did others he examined in late winter. Because the fat was in a lump rather than spread over each bird’s body, he speculated that it was being used for energy rather than for insulation. Given that these descriptions are for only a few birds, it is premature to conclude that this fat storage is common among roadrunners, but the reports suggest a good area for further research. Regardless of how the roadrunner survives winter’s cold and lack of food, the bird has proved that it is able to do so, and particularly in the northeastern parts of its range, where

34

The Real Roadrunner

it has been a permanent resident for more than fifty years. This trait speaks to the adaptability of the physiological processes and behaviors of this species. However, its tolerance of severe winter conditions has limits, which are discussed in chapter 8 in the context of possible northern extensions of the range of this species.

Surviving Predators No eyewitness accounts of a predator killing or disabling an adult roadrunner have been reported. In all reported predator attacks on an adult roadrunner, it has escaped by making swift maneuvers or seeking cover. The most common daytime attacks come from hawks, which dive at roadrunners in open areas. Upon spotting a hawk, the roadrunner crouches and freezes, with head turned sideways and upward to keep sight of the hawk. This posture is shown in the drawing at the beginning of this chapter. When the hawk dives, the roadrunner swiftly outmaneuvers the hawk with fast zigzag runs and leaps. Roadrunners have escaped attack by a red-tailed hawk, several northern harriers, and a Cooper’s hawk (Sutton 1977; Beal and Beal 1978; Douglas 1984; Meinzer 1993; author’s observation). Other animals observed to stalk or attack roadrunners during daytime include coyotes, bobcats, and domestic cats. No observations of snakes attacking adult roadrunners have been reported, except when parent roadrunners defend their nest from attack by a snake. In that case, the snake may strike back at the adult bird but soon retreats. The only confirmed record of a roadrunner in the stomach of another animal is that of a roadrunner in a red-tailed hawk’s stomach in West Texas (Van Tyne and Sutton 1937). However, this could have been a road-killed bird, since hawks will eat carrion. Observers recently reported circumstantial evidence of roadrunner predation by the endangered aplomado falcon in desert grassland habitat in Chihuahua, Mexico (Montoya, Zwank, and Cardenas 1997). Most students of roadrunner life agree that humans account for more roadrunner deaths than do daytime predators. The bird is often a traffic victim, despite its swiftness. And, as previously mentioned, the roadrunner is probably still hunted in some areas. Although no records exist, I suspect that most natural predation on adult roadrunners comes from nocturnal predators, such as owls, ringtail cats, foxes, weasels, and coyotes. The roadrunner’s nighttime lethargy associated with its temperature drop makes it vulnerable and unable to escape such predators. One cold night in the Sonoran Desert, my headlights caught a kit fox crossing the road with a roadrunner in its mouth. The bird was straight-asan-arrow rigid and immobile, as if in a stupor, rather than struggling or drooping in the fox’s mouth. Its propensity to seek a high, sheltered roost offers the roadrunner some protection from nocturnal predators. Wild adults that I watched in wooded areas would roost as high as twenty feet in oaks, cottonwoods, and other trees. Other roosting places have included within cholla cacti and on the tops of large barrel cacti. Roadrunners in urban areas roost on window ledges, in abandoned buildings, in shelves in open garages, and in chicken houses. My captive birds roosted on the highest perch available in their pen or in my house.

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Roadrunners require a sturdy tail support for roosting, for they back into a corner or against a tree branch with the tail held vertically upward. Two San Diego roadrunners roosted through a winter, each on a windowsill of a house. These birds must have been in the typical lethargic stupor, for they were not bothered by people or dogs being within inches on the other side of the glass (Abbott 1940). Predation on roadrunner eggs and nestlings probably accounts for more losses than does predation on adults. Nest predation and the parents’ responses to it are discussed in chapter 5.

4 Breeding Characteristics and Courtship

ow we will examine the events that take place during the breeding season of the roadrunner, when its daily life changes dramatically. In this chapter, I describe the courtship calls and displays that occur during this time. The following two chapters describe the events of nesting and how courtship unfolds during the breeding cycle, from egg laying through caring for the young.

N

Breeding Characteristics First, we will look at three characteristics of roadrunner breeding that make it successful: long breeding season; pair bond, with both parents sharing nesting duties; and courtship feeding before egg laying. These characteristics are described below, followed by descriptions of the courtship calls and displays.

Long Breeding Season For three-fourths of the year roadrunners are courting and nesting, from late winter into the fall. Because they don’t migrate, they can begin early and extend breeding into the fall in years when weather and food conditions are favorable. In good years they are able to have two and occasionally three successful nests. More than any other aspect of the roadrunner’s life, breeding behavior demonstrates the flexibility of the species in unpredictable environments.

The Real Roadrunner

38

Table 2 Breeding-season stages and approximate durations at a roadrunner nest

Days 1–5 6–10 11–15 16–20 21–25 26–30 31–35 36–40 41–45 46–50 51–55 56–60

Selecting nest site

Nest building

X

X X X X X X

X

X

Egg laying

Incubating eggs

X X

X X X X X X

X X

X X

Caring for nestlings 1–10 days

X X X X

Caring for nestlings 10–20 days

X X X X

Feeding fledgings

X X X X

Source: Whitson 1975; author’s unpublished field notes. Note: Table data are based on the following assumptions: A clutch contains four eggs, and an egg is laid every other day. The egg laid on day 6 would hatch by day 26 and fledge by day 45. The egg laid on day 15 would hatch by day 35 and fledge by day 54. The parents renest while nestlings are still in the first nest, as is typical of a second or third nesting in a season.

Several overlapping stages occur in the roadrunner’s breeding cycle, each with its own pattern of courtship and parental behavior. These stages and their approximate time frames for a nest with a four-egg clutch appear in table 2. While the parents are still feeding older nestlings and juveniles, they may start the process all over at a nearby nest site. Courtship calls, courtship displays, and mating are most frequent during nest building and egg laying. Courtship behavior tapers off while the pair care for eggs and young. As nestlings begin leaving the nest, courtship and mating attempts increase. This amorous activity may lead to a second or third nesting by the pair, except very late in the season. This long breeding season also maximizes the roadrunner’s potential to take advantage of abundant food supplies whenever they occur throughout the season. Although the peak egg-laying months reported from breeding bird surveys are April, May, and June, these data may be misleading, given that most breeding bird surveys do not extend beyond midsummer. In the Sonoran Desert of Arizona, roadrunner breeding peaks occur from mid-April to midJune and again from late July to mid-September. This latter peak follows the midsummer rains, when reptile food is abundant (Ohmart 1973). In South Texas, peak nesting extended through spring into July (Folse 1974; Folse and Arnold 1978).

Breeding Characteristics and Courtship

39

The earliest recorded date for egg laying is February 2, for a nest with four eggs found by James Cornett (1983) in Southern California (Riverside County). He noted that this early nesting occurred after a five-month period of higher-than-normal rainfall and slightly above-normal temperatures. This pair would have begun courtship and egg laying by late January. Other February dates for egg laying come from Kern and San Diego counties, California; Maricopa County, Arizona; Doña Ana County, New Mexico; and Zapata County, Texas (Cornell Laboratory of Ornithology, n.d.; Arizona Game and Fish Department, n.d.; author’s observations). The latest recorded date for eggs in a nest is October 10, reported by ornithologist Harry Oberholser (1974) for a nest in Texas, with no specific location given. Another late egg record comes from Duval County in the brush country of South Texas, for a nest with three eggs and an attending adult roadrunner at the nest on September 8. Several observations of nestlings in September come from Morrilton, Arkansas; Doña Ana County, New Mexico; and Brooks County, Texas (Sutton 1967; Cornell Laboratory of Ornithology, n.d.; author’s observations). A captive roadrunner pair nested three times during the year, with egg laying in June, July, and December at the Riverbanks Zoological Park in Columbia, South Carolina (Smith 1980). Yellow-billed cuckoos also have a long breeding season, lasting from April to September. Ornithologists William and Marian Hamilton (1965) suggest that this long season allows the species to take advantage of caterpillars and other insect food whenever it is plentiful, which is also likely the case for the roadrunner.

Pair Bond In some bird species the pair comes together only briefly for courtship and the female is left to care for the eggs and nestlings by herself. In other species, including the roadrunner, both parents care for the eggs and the young, and a long courtship period is common. Roadrunner courtship and mating continue during nesting, contributing to coordinated parental behavior and the ability to renest. This courtship promotes a pair bond that lasts throughout the breeding season and also may facilitate individual recognition of the mate and possibly mating for life. One of the mysteries of roadrunner breeding is whether or not these birds mate for life. It is known that the pair stay in the same area year-round, with each bird foraging in different parts of the territory when not breeding. They may even forage together at times during the nonbreeding season. By marking and later identifying individual birds over a two-year period in South Texas, Leon Folse (1974) confirmed that a breeding pair may stay together for at least two breeding seasons. Wild roadrunner pairs I watched stayed together throughout one breeding season, having as many as three nests together. However, since I didn’t mark these paired wild birds, I was unable to confirm that they were the same birds the next year. That roadrunner parents stay together for two breeding seasons could indicate a strong pair bond and mating for life. On the other hand, it could be that the same pair gets together

40

The Real Roadrunner

each year simply because they forage in the same area and are near each other when the next breeding season starts. Banding data show that roadrunners travel over only a small area during their lifetime (Patuxent Wildlife Research Center Bird Banding Lab, n.d.). I saw evidence at several nests that male roadrunners may bond with more than one female. At West Texas nests, male roadrunners exhibited courtship behavior to various roadrunner models placed near the nest (Whitson 1971, 1975).

Courtship and Nuptial Feeding A prominent feature of the male’s mating behavior is his feeding of the female. This feeding may nourish her before, during, and after egg laying. Courtship feeding in birds occurs primarily in species that share parental duties (Lack 1940). Courtship feeding occurs in many other cuckoos, as well as in other bird families, although it is only in cuckoos that the food exchange happens during or after, rather than before, mating. About three-fourths of the matings I witnessed involved food. The female usually received the male when he offered a lizard, a snake, or a mouse. Most of the unsuccessful matings occurred when the male offered insects, twigs, or nothing. Wyman Meinzer (1993) also observed that females rebuffed males that brought insects rather than larger prey. On the other hand, I sometimes observed mating with no courtship food. Such matings usually took place after the eggs had hatched. Nuptial feeding in birds may be necessary to provide the female with sufficient nourishment for egg laying. In some bird species, breeding coincides with times when the female has enough food to form eggs (Perrins 1970). This could be the case for the roadrunner, because most courtship food is offered to females before and during egg laying. Feeding the female may also ensure that nesting occurs at a time in the season when abundant food will be available for the nestlings. This timing is especially important in a habitat where the food supply is variable during the season or from year to year. Courtship feeding may help synchronize nesting with times of sufficient food for the nestlings. Because the courtship and nuptial food contains more vertebrates than does the female’s normal diet, it resembles a nestling diet rather than an adult diet. Unfortunately, there are no reported observations of courtship displays or courtship feeding in the roadrunner’s eastern or northern habitats to tell us what food the male roadrunner uses to woo the female in these habitats.

Calls and Displays of Courtship Courtship Calls When Far Apart Courtship displays of many bird species occur in the air and are visible to their distant mate. Such aerial acrobatics are not possible for the terrestrial roadrunner. My research (Whitson 1971, 1975) on roadrunner vocalizations indicates that calls play an important role in the courtship of roadrunners. Their far-reaching courtship calls may be the best way for widely scattered individuals to locate and attract each other. When separated, the pair’s

Breeding Characteristics and Courtship

41

Figure 7. Head movements of a six-note coo call. Each drawing represents the production of one note. The solid line depicts the head’s position just before the note begins; the dotted line, the head’s position at the end of the note.

loud bill-clacking sound and cooing and barking calls may keep them in touch and enable them to find one another. The mournful coo of the breeding male is the one that is the most familiar to those who live in roadrunner country. Some people mistake it for the sound of a cooing dove, which it somewhat resembles. This call is common during the breeding season, but especially so in the early spring. The male usually calls from the top of a bush, a tree, or some other high perch for up to several hours at a time. The coo typically consists of five notes but ranges from three to eight notes. Individual males usually have distinctive coos but may vary the number of notes in their coo at different times. The greatest individual variation I taped was for a wild male whose coo ranged from four to eight notes. Specific head movements and throat bulging go along with the coo and are probably essential to producing the sound (figure 7). The bird lowers its head, with its crest raised and its orange patch showing. As each note is made, it moves its bill away from its chest, then moves it back to the chest before the next note begins. The first note is always the shortest and the last note the longest. A typical five-note call sounds like this: co-coo-coocoo-cooooooo. It can be heard by humans from several hundred feet away from the bird. The call appears to attract the female and to maintain contact when the pair are foraging separately or are feeding widely scattered fledglings. It also may be used to declare the breeding territory. Two birds are commonly heard giving alternating coos at a great distance from one another. On the few occasions when I have witnessed territorial fights, both birds gave the coo before, during, and long after the fight. Some observers have reported females also giving the coo, although I have never observed this in wild or captive pairs. The yellow-billed cuckoo also has a many-noted coo, with head motions similar to those that accompany the roadrunner’s coo. Likewise, the yellow-billed cuckoo gives its call for long periods from a high perch and varies the number of notes from time to time.

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The Real Roadrunner

The bark is another call that is used often when a roadrunner pair are separated. The call is a series of two to twenty-two short, rapidly repeating notes that resemble canine barking or yelping. I have heard both wild and captive females give this call. The bird’s bill is open as its body jerks slightly with each bark note. The female gives this call often in response to the male’s distant coo early in the season and later while she is incubating the eggs or brooding the nestlings. She also barks in response to distant barks, suggesting that the bark could be a territorial call as well. I do not know if wild males also bark, but none of the males I raised gave this call. The bark is the most frequent call that humans hear during the nest-building period. The pair often exchange bill clacks. The clack is made by the upper and lower bills rapidly snapping together and is accompanied by a throaty, whining sound. It is sometimes referred to as a rattle. The clack, which is highly variable in loudness, consists of five notes on average and ranges from one to twelve notes. Both sexes give the clack, and nestlings also snap their bills together, producing a soft, hollow version of the clack. The parental pair exchange clacks when separated early in the breeding season and when temporarily separated during foraging, suggesting that clacks may be used to maintain contact. Throughout the year, the roadrunner clacks when it is startled or agitated.

Courtship Displays and Calls When Close Together In many bird species the plumage of the male is more colorful than that of the female, a characteristic that is thought to help the male attract a mate. Despite the dull plumage of the male roadrunner, he uses what physical features he has to good advantage as he strives to attract and court a mate. Although small, the bright orange patch on the male’s head is eye-catching, especially from a side or back view (plates 2 and 3). Likewise, the movements of the crest and the long tail, tipped with conspicuous white spots, are attention getters. These head and tail movements, combined with coos and whirrs in persistent, coordinated displays in front of the female, are quite impressive. While near the female, the male turns and runs from her, throwing his wings upward until they meet above the back of his body (fig. 8a). He then quickly lowers his wings, with a downward-inward wing motion (fig. 8b) that results in a loud, popping sound. This display usually continues for four to five wing-raising cycles, with the bird running away from the female the entire time. The male closes his tail and holds it forward over his back during this display, which I call the prance (fig. 8c). Because his back is to the female, this tail position shows off the prominent white tail tips, normally not seen. The bright orange patch behind the eye is also uncovered during this display. Although the prance is part of courtship, it may also be used as a territorial display between males. Some species of male hummingbirds exhibit an aerial wing popping, which is thought to attract females. The spruce grouse has a wing-snapping display on the ground that produces a popping noise.

Breeding Characteristics and Courtship

a

b

43

c

Figure 8. Prance display of male: a, running with lifted wings and tail, back to female; b, lowering his wings to his body as he continues running; c, side viewof b.

Tail wagging is a male display, although on a few rare occasions I observed tail-wagging behavior by females. This is the most common male courtship behavior and usually involves courtship food (plate 9). The male wags his tail from side to side, as he bows and then slowly lifts his head. The head and tail movements are synchronized so that when the head is up, the tail is closed and swings from side to side (fig. 9a), and when the head bows, the tail fans (fig. 9b). The tail is stationary for a moment during the bow, then begins swinging from side to side as the head is lifted (fig. 9c). Tail fanning reveals the white tips of the tail feathers, and the bright orange patch is visible as the male bows his head in front of the female. A whirring call is coordinated with the tail and head motions. The call is a steady stream of soft, low-frequency putt sounds, regularly interspersed with a slow, whirring sound. The effect can best be described as putt-putt-putt-putt-whirrrrr-putt-putt-putt-whirrrrr. The male gives this call while he bows and raises his head. The putts of the call occur while the head is erect or being lowered, and the whirrrr as the head lifts from the bow. The call is very soft and can be heard by humans only within a few feet of the bird. I was able to tape-record this call only because one of my hand-reared males imprinted to humans and courted me vigorously during the breeding season. The male usually begins wagging his tail about thirty yards from the female as he walks or runs toward her. As he draws near, he stops often to repeat the tail wag, and the

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a

b c Figure 9. Tail-wag display: a, lateral extremes of tail motion when bird’s head is erect; b, tail fanned and stationary, with bird’s head bowed; c, tail fanned and starting to wag as the bird’s head lifts from the bow.

whirr call becomes louder the closer he gets to her. Often he will coo during one of these stops. This coo is a softer and shorter one than that given when the birds are separated. The striped cuckoo also uses a tail-wagging display during courtship, but this bird’s display does not include head bowing. Another species with prominent white tail tips, the Inca dove, has a courtship display with a raised, fanned tail. The vertical flick is a rapid, upward tail movement by the female roadrunner in response to the tail wag of the male. The female gives the vertical flick, along with a bill clack, only at those times when she allows the male to mount. As the male approaches and begins his mounting jump, she turns her back, flicks her tail upward, then lowers her tail as he mounts. The female yellow-billed cuckoo has a similar tail-up posture as part of her mating display. When the female is not receptive to the tail-wagging male, she flicks her tail to the side, rather than upward, and drops it as she moves away. Both sexes give a similar sideways tail flick throughout the year when approached closely by another roadrunner or when they encounter a strange object.

Breeding Characteristics and Courtship

Figure 10. Female taking lizard from male at end of mating.

45

Figure 11. Mating with no food exchange. Note the upward-reaching position of the female’s head.

As the female moves away, the male may circle her while wagging his tail intensely or he may leave the area. He either eats the courtship food himself or takes it to the nestlings if the eggs have hatched.

Mating Mating begins as the female lowers her breast and wings while the male jumps and lands on her. He stands on her upper wings and rapidly stomps his feet. The female supports herself on her chest, upright legs, and outspread wings (plate 10). Just before mating is over, the female grabs the courtship food from the male’s bill (fig. 10). My movies of wild birds mating show that even when no courtship food is exchanged, the female still moves her head upward at the end of the mating (fig. 11). A wild female once provided me an amusing moment. She jumped out from under the male before mating was over and quickly grabbed the lizard he was holding. The male refused to release the lizard, resulting in a vigorous tug-of-war, with the male holding the lizard’s head and the female the tail. The male finally pulled the lizard away from the female and ate it himself. I hoped that the lizard was dead during this tug-of-war. After mating, the pair give a coordinated tail and head-flicking display. The male circles the female, which remains in one place and gradually turns her body to face the circling male. The male stops every few steps, bows his head, lowers his wings slightly, gives a single soft coo, and then quickly flicks his head, wings, and tail upward. The female gives this flicking display also, sometimes in unison with the male (fig. 12). The female eats the courtship food until the eggs hatch, when she feeds it to a nestling instead. After the young fledge, the female eats the courtship food rather than giving it to the fledgling.

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Figure 12. Mutual flick display by both sexes after mating.

Other Interactions between the Sexes While foraging together, one bird may approach its mate with a stick or a blade of grass in its bill and drop it or transfer it to the mate’s bill. This occurs in early interactions between the pair as well as later during nest site selection and nest building. The yellow-billed cuckoo also offers sticks as part of courtship, as do many other birds. A soft, whining call is often, but not always, associated with stick giving. This call is a long, low-frequency whining note given by both sexes. The note is often repeated for several minutes, with the bird in a crouched position on the ground or on a branch. Its head moves downward and side to side while giving the call (fig. 13). The growl call is a three- or four-note, low-frequency growling sound, made as the bird’s throat bulges noticeably with each note. Both sexes give this call, especially as they forage close together. Pairs also use the growl while building a nest, feeding the nestlings, and leading the nestlings from the nest. The bark and the clack calls are also heard frequently while the pair is foraging together and building the nest. Some of these calls appear to be in response to distant calls of other roadrunners, rather than directed toward the mate. What appear to be aggressive displays between the sexes occasionally occur before the nest is built. These displays usually occur while one bird is chasing another. The pursuing

Breeding Characteristics and Courtship

47

Figure 13. Female giving the whine call from a perch.

bird runs at the other in an attack posture, although physical contact is not made. This posture also occurs in territorial fighting, when one roadrunner invades another’s territory.

The Role of the Coo Given that the coo is the common call during the breeding season, I examined its role in courtship by playing a coo recording to wild birds and observing their reactions. Playing the call alone removes the possibility that the bird is responding to the sight of a visual display made by the calling bird. To my surprise, wild roadrunners responded to a tape of the coo not only with calls but also with some of the visual displays of roadrunner courtship. Nearly 90 percent of the more than two hundred roadrunners tested approached the source of the coo call. Nearly half of the birds cooed in response, and another 12 percent barked. About 10 percent of the birds tail-wagged or pranced, though no other roadrunner was present. Six birds approached with a lizard or an insect in their bill as they tail-wagged. Several approached without food, then left the area, soon returning with a lizard or an insect in their bills. In these coo tests I did not know the sex or breeding status of the birds that responded, and some of them could have been unpaired or juvenile. I also played the coo tapes to paired wild birds whose sex and phase of breeding I knew. The male responses to the coo tape included cooing, prancing, and tail wagging, while the female responses included barking, whining, stick presentation, and sideways and upward tail flicking. All of the male responses increased as the nesting cycle progressed from pre–nest building to the time the nestlings fledged. However, the females were responsive only before and during nest building.

5 Getting Together, Nest Building, and Incubating

fter watching roadrunner parents at several nests, I realized that successful nesting requires not only nesting skills but also efficient coordination of duties between the sexes. The first challenges that potential parents face is to come together as a united pair after foraging separately during the winter. Once together, they must agree on a safe place to build their nest. After egg laying starts, they face a myriad of hardships and irritations, including egg-eating snakes, ants in the nest, and temperature extremes. On top of all this, they are unwelcome neighbors in the local bird community. Other nearby nesting bird species repeatedly dive at roadrunners to keep them away from their own eggs and nestlings, which roadrunners are prone to eat or feed to their own young. In this chapter and the next we will see how roadrunners overcome the odds to get together, mate, and successfully raise their young. Nesting data, including nest heights, clutch size, and types of nest structures, came from the North American Nest Record Card Program (Cornell Laboratory of Ornithology, n.d.); Arizona nest records (Arizona Game and Fish Breeding Bird Survey, n.d.); and the author’s field notes from twelve nests studied in depth and sixteen other nests studied briefly. These nests were in Oklahoma, New Mexico, and West Texas.

A

Getting Together In late winter or early spring the first signs of male breeding behavior are frequent coos throughout the day. The male appears bent on declaring his intentions throughout his territory, for after cooing for a while in one spot, he moves a few hundred feet away and begins

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again. Exactly what his intentions are at this time is not clear, for besides the time for courting, this is the time for declaring and defending the breeding territory. Often a distant roadrunner coos or barks during or immediately after the first bird’s coo call. Occasionally, two distant birds appear to be exchanging bark calls. Direct encounters with other roadrunners in the area bring forth behavior ranging from intense tail wagging with courtship food to aggressive chases and attacks. During this time it is difficult to tell territorial behavior from early courtship behavior. One bird will chase the other or vice versa, often for several hours. Fast running is interspersed with low, gliding flights, frequent rest stops, tail wagging, prancing, and cooing. Occasionally, the pursuing bird attacks with its wings spread and its tail raised and fanned. Most likely these chases are between males, as well as between male and female. Since this attack posture is like that of territorial aggression between males, some chasing may be territorial rather than courtship behavior. During my field observations, I saw actual fights between roadrunners only four times. The drawing at the beginning of this chapter shows two birds in a territorial encounter. The first sign that the pair is established is when they forage together for most of the day. However, they still roost separately for a few days. Each morning, separated once again, the male’s coos and the female’s responding barks help them get together for the day. They often exchange calls, especially a soft growl call, as they move in the same general direction. Soon the pair starts roosting close together, often in the same bush or tree. Nest-building behavior begins with small sticks being passed back and forth between the pair. One member of the pair whines while perched on a branch or other aboveground spot, followed by the other bird searching for a stick to give to it. The whining bird usually drops the stick and begins foraging again. The male will sometimes prance or tail-wag near the female during this time, but mating doesn’t occur at first.

Selecting Safe Nest Sites Soon after the birds start giving sticks to one another, they begin looking for the right nest site, which appears to be a joint effort. The pair may be testing sites as they move together into shrubs, trees, or other high perches. The female often whines as the two perch close together or side by side in a tree or shrub, prompting the male to leave and return with a stick for her. She may begin nest-building motions with the stick or drop it, in which case the pair continue to search for the perfect site. Male courtship displays and female whining occur in the midst of this search. Mating occasionally occurs but is not frequent until the pair begins to build the nest. We don’t know what prompts the birds to choose a particular nesting site, but looking at common features at many nest sites helps define a few of the bird’s requirements. I compiled unpublished nesting information from the North American nest card record program (Cornell Laboratory of Ornithology, n.d.) and the Arizona Breeding Bird Atlas (Arizona Game and Fish Department, n.d.), along with published nesting records and my nesting observations.

Plate 1. Adult roadrunner in Chihuahuan Desert of West Texas.

Plate 2. Male roadrunner with crest erect and colored skin patch displayed; captive bird of Oklahoma origin.

Plate 3. Male roadrunner with crest partially erect and colored skin patch displayed; captive bird of Oklahoma origin.

Plate 4. Recently fledged roadrunner, about twenty-five days old; Chihuahuan Desert of West Texas.

Plate 5. Roadrunner starting to swallow Texas horned lizard. Photo provided by Dr. Wade Sherbrooke.

Plate 6. Roadrunner swallowing Texas horned lizard. Photo provided by Dr. Wade Sherbrooke.

Plate 7. Juvenile roadrunner absorbing heat from the sun on the bare black patch on its back.

Plate 8. Roadrunner sunning during the heat of the day. The function of this sunning behavior is not known but possibly is to rid the bird of feather parasites.

Plate 9. Captive male roadrunner holding Chihuahuan spotted whiptail lizard during the tail-wagging display of courtship. Photo provided by Dr. Wade Sherbrooke.

Plate 10. Mating of roadrunners; Chihuahuan Desert of West Texas.

Plate 11. Roadrunner nest with five eggs under attack by bull snake; Chihuahuan Desert of West Texas.

Plate 12. Roadrunner nest with ten eggs, six feet high in a honey mesquite tree in West Texas.

Plate 13. Female roadrunner incubating the ten eggs at the nest shown in plate 12.

Plate 14. Coiled western diamondback rattlesnake, with its head hidden under its coil, after an attack by a wild roadrunner.

Plate. 15. Nesting adult roadrunner attacking a taxidermic roadrunner mount placed near its nest in West Texas.

Plate 16. Roadrunner nest with four chicks, four days old and younger; Chihuahuan Desert of West Texas.

Plate 17. Twenty-five-day-old juvenile in immobile, erect posture in response to nearby human; West Texas.

Plate 18. Month-old juvenile begging for food from its parent.

Plate 19. Head of four-to-five-day-old chick.

Plate 20. Foot and wing of four-to-five-day-old chick.

Plate 21. Fifteen-day-old roadrunner nestling; Chihuahuan Desert of West Texas.

Plate 22. Twenty-three-day-old juvenile, panting and with drooped wings to allow body heat to escape.

Plate 23. Two-month-old juvenile in West Texas.

Plate 24. Seventy-five-day-old juvenile, with still-blotched juvenile mouth and eye ring of adult.

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These 254 nesting records come from ten states, but the majority are from southern parts of Arizona, New Mexico, California, and Texas. Most roadrunners select an aboveground “structure” that will provide some cover, such as a cactus, a bush, a tree, or an artificial structure. Only two of these nest records were on the ground. The average nest height was 6.8 feet (for the 190 nests for which nest height was recorded). The highest nest reported was 25 feet aboveground in an evergreen tree in Colorado. The only ground-cuckoo nesting record is for a rufous-vented ground-cuckoo nest, located more than 8 feet high (P. Roth 1981). In upland brush habitat in South Texas, Folse (1974) reported an average nest height of 4.6 feet for twenty-eight nests, most of which were in bushes. Ohmart (1973) reported an average nest height of 4.3 feet for sixteen nests in the Sonoran Desert of southern Arizona, most of which were in cholla cacti. An average nest height of 4.7 feet was reported for twelve nests in the Chihuahuan Desert of West Texas (J. M. Hughes 1996a). Safety from predators is an obvious reason that this ground-dwelling bird takes to the heights when it is time to nest. The eggs and newly hatched nestlings are especially vulnerable to predators. Until nestlings are about twelve days old, they are not developed enough to walk well, unlike the chicks of many ground-nesting birds, such as killdeer or quail, which can run immediately after hatching. Typically, the parents chose progressively higher nest sites for the second and third nests of the season, but not always. A New Mexico pair I watched reared their first clutch in a nest seven feet high in the heart of a yucca and their second successful clutch in a nest ten feet high in a nearby tree, but their third successful nest of the season was only three feet high in a small bush. Roadrunners seem to prefer nest sites in the middle of or on the edge of open areas. By having an open area near the nest, a parent can keep a watchful eye on the eggs or nestlings while it forages, suns, preens, or mates. Roadrunners in upland brush habitat in South Texas nested in small isolated bushes, even through clumps of bushes and trees were scattered throughout the area (Folse 1974). Several other nest site features may be important to roadrunners. Most nests are positioned so that the nest surface is shaded or partially shaded during the hottest times of the day. In West Texas, Janice Hughes (1996a) observed that nest bushes were generally less dense on one side than on the other and had a greater distance to the lowest branch than adjacent bushes did. She suggested that these features might facilitate the roadrunner parent’s entering and leaving the nest bush. Roadrunner parents typically approach a nest from the ground by climbing upward from branch to branch and leave the nest by gliding downward to the ground. Roadrunner parents do not appear choosy about the type of support structure, as long as it provides an adequate base for their bulky, loosely built nests. They most often nest in cacti in Sonoran Desert habitats and in woody plants in South Texas brush habitats. Prickly pear and cholla are the typical cacti used for nesting. A variety of trees and shrubs have served as nest sites, reflecting what is available in the habitat. Types of woody plants used for nesting

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include mesquite, white brush, huisache, colima, juniper, cedar, salt cedar, cottonwood, willow, elm, olive, Russian olive, pine, oak, hackberry, and ebony. The birds also place nests within or atop large brush piles in South Texas brush habitats (author’s observation) and in crevices in sandstone cliffs in California (W. L. Dawson 1923). The variety of artificial nesting structures that the roadrunner uses shows its resourcefulness. These have included oil derricks, old farm machinery, old plane wheel wells, interiors of abandoned cars, window ledges, and shelves in open garages or deserted buildings.

Building the Nest After the nest site is selected, the female does the building while the male brings her the twigs. If he dallies, she lowers and wobbles her head from side to side as she gives the whine call. As the nest takes shape, she sometimes sits in it and wiggles her body into the center. The nest is built within a day or two, although the pair keeps adding twigs and grass until the young leave the nest. Sometimes the site doesn’t seem to please the female, for I have seen several females leave the nest midway through building and begin the site search once more, with the male following. The nest is usually eleven to twelve inches wide and six to eight inches high, although it may vary in width from seven to seventeen inches (Sutton 1940; J. M. Hughes 1996a). It consists of a bulky bottom layer of loosely laid, large twigs and a soft, inner bowl of small stems and leaves. The pair enhances the nest lining with soft material, including feathers, snake skin, and other debris. One California woman told me that she provided cotton balls, which were used as nest lining by the roadrunner pair nesting in her yard. Naturalist Joseph Grinnell (1893) reported that roadrunner nests in California were lined with small bits of dry horse manure. At West Texas nest sites I observed, three-fourths of the successful matings took place while the pairs were building nests and laying eggs. Courtship calling, tail wagging, courtship feeding, and mating occurred more frequently during nest building than at any other time, except for the coo, which was more frequent before nest building. The bark call was common during nest building, and the female often exchanged barks with a distant bird. Both sexes gave the growl call as they foraged together.

Laying the Eggs Egg laying begins before the nest is in its final form and occurs at any time during the day. By checking nest contents during the egg-laying period, I found that eggs are laid quickly, often during only a twenty-minute sitting period. Sometimes an egg is laid every day, but more commonly every other day. The time between eggs can be as long as nine days (Folse 1974). Eggs are off-white, with a chalky surface film, and are oval to elliptical in shape, with some shape variation within a clutch. The average size is 1.6 inches long by 1.2 inches wide. Clutch size varies from two to twelve eggs, with a clutch of four eggs being the most common. Plate 12 shows a ten-egg clutch at a nest I watched in West Texas. After finding a

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ten-egg nest where two chicks hatched the same day, George Miksch Sutton (1940) speculated that several females may have been responsible. I have seen more than two adults around a nest but have never found evidence that more than one female had laid eggs in a nest. The smooth-billed ani is a communal nester, with two or more females laying in the same nest (Skutch 1966). Roadrunner parents may accept eggs other than their own in their nest. A quail’s egg once appeared in a roadrunner nest in Arizona (Arizona Game and Fish Department, n.d.). A field test at a wild roadrunner nest found that the incubating parent would accept eggs different in size, shape, weight, color, and number from its own (Eltaher 1980). The bird incubated dummy eggs that were up to four times larger and four times smaller than its own egg. It also accepted increases and decreases in the number of eggs (double the number and half the number) and dummy eggs (wood, glass, plastic, spherical, colored, and chicken eggs). Little evidence exists that the roadrunner shares a brood parasitism habit with its cuckoo relatives. Only two instances of a roadrunner egg occurring in another bird’s active nest have been recorded. One roadrunner egg was found in a common raven nest along with three raven eggs in California (Pemberton 1925). Because Pemberton collected the eggs, we do not know if the raven would have reared the young roadrunner. A roadrunner egg occurred in a northern mockingbird nest along with one mockingbird egg in Texas, with no details given on subsequent events at the nest (Oberholser 1974). Courtship displays and calls are less frequent after egg laying starts than during nest building. The female’s bark, the most common call, is usually heard from the nest. The incubating bird and its nearby mate often exchange clacks. Mating is frequent, usually occurring just below the nest site or within a few feet of it. As the male approaches the nest, he will give the coo or clack and begin wagging his tail as he nears the nest site, often with a lizard in his bill. The female jumps or glides down to join him in the mating act.

Incubating the Eggs Roadrunner eggs hatch in nineteen to twenty days in the wild, although in the controlled conditions of a laboratory incubator they have hatched sooner. Artificial incubation records include one egg hatched in seventeen days (Smith 1980); four eggs hatched in seventeen to eighteen days (Calder 1967a); one hatched in eighteen and one-half days (Muller 1971); and another hatched in eighteen and one-half days (author’s observations). Incubation starts as soon as the first egg is laid, resulting in staggered hatching dates and various-sized nestlings. Having nestlings of different ages may ensure that at least some of them survive predator attacks or food shortages. Now around-the-clock sitting begins, with parents sharing the duties of incubation. During the breeding season, both sexes develop an unfeathered brood patch on the abdomen, which allows as much body heat as possible to transfer to the eggs. The blood vessels in this area increase in size, and the skin becomes thicker and more fluid-filled than normal.

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The parents alternate shifts during the day, but the male always spends the night on the nest (Ohmart 1973; Vehrencamp 1982; author’s observations). In nests I watched, females incubated eggs slightly more during the day than did males. Sandra Vehrencamp (1982) reported females and males incubating about the same amount of time during the daylight hours. In the morning the female does not relieve the male from his duty until one or two hours after sunrise. This gives her time to sunbathe and feed before beginning her morning shift, which might last for two to four hours. During his morning break, the male suns, feeds, and sometimes announces his presence to neighboring roadrunners by cooing from various high perches in the breeding territory. Occasionally, the female will bark or clack from the nest during or after the male’s episodes of cooing. By early afternoon the male returns to relieve the female for a several-hour stint; later the female returns for the midafternoon to early-evening shift. Shortly before dusk the male returns to begin nighttime duties. Sometimes he brings a lizard and begins cooing and tail wagging as he nears the nest. In response, the female clacks and jumps from the nest to meet him, and the pair mates on the ground near the nest. After eating the lizard, the female begins the trek to her roosting spot in another part of the territory. Along the way she preens, dust-bathes, and forages. While on the nest, a parent does not sit idly but busily attends to such housekeeping duties as regularly rolling the eggs. On the average, I saw eggs being rolled about every fifteen minutes by each parent. Along with egg rolling, the parent will often fiddle with the nest twigs and lining, pulling a twig out of the nest and moving it to a new spot or dropping it. Another important duty is keeping the nest clean and free from pests. Ants, mites, and other small creatures that invade the nest are quickly gobbled up when within the parent’s bill range. During long shifts, a parent may leave the nest for several minutes to forage and drink nearby, but it rarely moves out of sight of the nest. Often it will bring back grass or small twigs to add to the nest. The amount of time the eggs are left alone seems to vary little with the weather. I watched two nests with four eggs each in the same area in West Texas—one during an unusually cold, wet March, and the other during a hot, dry July summer. The summer parents left the eggs alone only slightly more than did the spring birds. Before the eggs hatch, the nest is left alone for no more than 5 percent of the day. After several eggs are hatched, the parents spend longer periods away from the nest than they do before hatching.

The Male’s Night Shift There are only a few other bird species in which the male spends the night on the nest. Groove-billed ani and smooth-billed ani males also take the night shift, as do male woodpeckers. This behavior may be more common in birds than is currently known. Sandra Vehrencamp (1982) proposed a good reason for the male rather than the female being on the nest at night, and it is related to the way roadrunners save energy. Ordinarily, adult roadrunners lower their nighttime temperature to avoid expending energy (see chapter 3). By absorbing

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heat from the next morning’s sun, they warm up quickly without using food energy. But because eggs must be kept constantly warm to properly develop and hatch, the incubating parent needs to maintain a normal body temperature at night. To determine the incubating male’s nighttime temperature, Vehrencamp (1982) attached temperature-sensitive radio transmitters to wild-nesting roadrunner pairs and to non-nesting males. She found that at night incubating males maintained their body temperatures about 10°F higher than did females and non-nesting males. That the male would be the one to use the energy needed to maintain a high nighttime temperature makes sense, for he is in better physical condition during incubation. An incubating male has conspicuous fat deposits and is heavier than a non-nesting male (Vehrencamp 1982). The female uses energy to lay the eggs and has very little fat reserve left after egg laying. By saving energy during the night, the female can rebuild her weight for the tasks of feeding the young and laying eggs for the next nest. Another benefit of the male’s taking the nighttime vigil is that it allows the female to roost in a safe spot away from the nest. Nighttime predators, such as ringtail cats and foxes, could locate the nest from the odors emanating from it.

Defending the Nest from Predators The only predators that have been documented at roadrunner nests are egg-eating snakes, such as rat snakes, coachwhips, and bull snakes. Other probable nest predators include coyotes, foxes, raccoons, skunks, bobcats, ringtail cats, and domestic cats. The amount of predation on roadrunner eggs and nestlings varies greatly from year to year and place to place. Robert Ohmart (1973) reported no nest predation at the nests he watched in the Sonoran Desert of Arizona. There was some predation at the ten nests I watched in Chihuahuan Desert areas of West Texas and New Mexico, but three-fourths of the eggs produced young that fledged. However, only slightly more than a fourth of the eggs produced young that fledged in Leon Folse’s South Texas nesting study (1974). Like most parent birds, the roadrunner will defend its eggs and nestlings at the risk of its own life. One of my most vivid roadrunner memories is that of a female attacking a very large bull snake in the thorny shrub that held her nest (plate 11). The snake began swallowing the first of the five eggs as the female attempted to jab at it from nearby branches. Without much room to maneuver, she was not able to kill the snake, but she finally chased it away after it had eaten one egg. The snake must have come back later, for the rest of the eggs were gone the next day. The pair abandoned this three-foot-high nest and built a new nest six feet high in a nearby honey mesquite. Besides doubling the height of the new nest, the pair also doubled the clutch size from five to ten eggs (plate 12). A parent is sitting on these ten eggs in plate 13. Six of these eggs hatched, but the rest disappeared, possible eaten by the same bull snake that attacked the birds’ first nest. Other accounts have been recorded of a parent defending its nest from a snake much larger than it could possibly eat. To determine if a roadrunner parent would defend its nest

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against a rattlesnake, I placed a three-foot western diamondback near a nest to observe the encounter. Quite a battle followed, with the roadrunner using rapid, jabbing motions to peck the snake’s head. It always avoided the snake’s strike by quickly leaping upward or by using its wings to divert the snake’s strike. As the bird ran toward the snake, it held its wings fanned and at right angles to its body, resembling the position of a bird during territorial fighting. At times the bird quickly flashed its wings, similar to the wing-flashing during foraging. The fight was over in a matter of minutes, for the snake coiled up and put its head under the coil (plate 14). This position prevented the roadrunner from pecking the snake’s head, the only way a roadrunner can kill a snake. When the snake refused to uncoil, the roadrunner stopped striking at the body and returned to its nesting duties. Even when picked up afterward by a human, the snake stayed in this rigid, coiled position for some time. Wade Sherbrooke witnessed a similar defensive reaction by a western diamondback when it was attacked by a roadrunner (pers. comm.). A parent will also protect nestlings from predators or humans by trying to lure them away from the nest. Whenever I got too near some nests, the parent would jump to the ground, run in a crouched position, and circle the nest area until I moved out of sight. A roadrunner observer in California reported a ploy used by a roadrunner with five nestlings to protect. This bird simulated a broken leg and dragged itself away from the nest, sometimes falling over on its side (Pemberton 1916). Nesting roadrunners will also protect the nest from others of their species. During experiments with roadrunner models near breeding pairs, I found that some painted wooden models and taxidermy mounts were vigorously attacked and torn to shreds when placed near the nesting site. Both sexes would attack these models (plate 15).

6 Caring for Nestlings and Fledglings

ince roadrunners begin incubating with the first egg rather than waiting until a full clutch is laid, the eggs hatch sequentially rather than at the same time. Nests with large clutches may have older, feathered nestlings about to fledge alongside a younger, unfeathered chick. Even after all the eggs hatch, parents spend most of the time sitting on the nest. For about twelve days after the first egg hatches, either the male or female is at the nest for most of the day.

S

Caring for Nestlings Brooding Nestlings Why this prolonged brooding after the eggs hatch? Because a newly hatched roadrunner cannot control its temperature and will rapidly lose or gain heat through its bare, black skin, the brooding parent is crucial to its survival. During early spring nesting a parent may need to brood the nestlings to keep them warm and dry. On hot, sunny days a parent needs to stand over the nest, shading the young during the times that the sun’s heat could raise their body temperatures to dangerously high levels. Because of the staggered hatching schedule at a nest, parents may be brooding the younger, naked nestlings along with older, feathered nestlings that no longer need brooding. It is amusing to watch a nest in which a parent is snuggled on top of younger nestlings, while older nestlings teeter on the edge of the nest or on a nearby branch. When one of these older chicks gets trapped under the brooding parent, it wiggles out, fluffs its feathers, and clacks its bill, as if protesting being held under the hot body of the parent.

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After all nestlings are at least partially feathered, the parents spend more time away from the nest, foraging for additional food for the rapidly growing young. After older nestlings fledge, the parents divide their time between feeding the nestlings and feeding the fledglings.

Feeding Nestlings “Ravenous” is the best word to describe nestling roadrunners. Several gaping, reddish pink mouths confront a parent each time it brings food to the nest (plate 16). Nestlings that cannot yet stand wave their gaping bills in front of the parent, while older chicks stand gaping and rapidly flutter their wings. Begging nestlings make a continuous, loud sound that is part buzz and part whine. The nestling diet consists primarily of reptiles and insects (see chapter 2). A parent usually brings one food item at a time, leaving one or more nestlings still hungry and begging persistently. It is no wonder that a parent sometimes brings two pieces of food at once, such as a lizard and a grasshopper or two lizards. The amount of daily food a nestling eats usually weighs half or more of its own weight. Even a newly hatched chick may be fed a small lizard. I have seen as many as five lizards fed to one nestling in a day. A parent does not beat vertebrate food before giving it to a nestling, in contrast to the prolonged beating it does before it eats vertebrate food itself. Evidently the nestling’s digestive system is able to handle this animal food, for nestlings don’t regurgitate pellets with bones, fur, feathers, scales, and other animal parts (see chapter 2). A parent usually feeds whichever nestling is begging most vigorously, but sometimes a parent ignores an aggressive nestling and gives the food to another. A parent often will keep its grip on the food for from twenty seconds to two minutes, and as the nestling swallows more of the food, it appears to swallow the parent’s bill as well. An adult sometimes makes slight pumping movements with its head while holding the food, as if stuffing it into the nestling’s throat. When a parent feeds a long snake, it holds the rear of the snake up while the nestling swallows the head end first. A parent rarely holds food while a ten-day-old or older nestling is swallowing.

Food Shortages In times of drought, cold springs, or fall nests, a good food supply for nestlings may be lacking, and older nestlings might outcompete younger ones for food. Robert Ohmart (1989) reported observing a roadrunner parent eating a weak, lethargic nestling at two nests in Arizona. A young roadrunner was recently reported in the stomach contents of an adult roadrunner in Texas, although whether it was the adult’s own young is unknown (Kazmaier, Ruthven, and Synatzske 1999). It has been suggested that older nestlings may eat their younger siblings, but there are no observations to support this. With its weak neck muscles, a nestling would have a hard

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time picking up one of its siblings and swallowing it. Young roadrunners that I observed did not pick up large food items for themselves until they were about twenty-five days old.

Keeping the Nest Clean A clean nest is important to nestling and brooding parent survival. If parents do not remove nestling wastes promptly, strong odors may attract predators. The urine and excrement of nestling birds are packaged together in mucous sacs called fecal sacs, which can easily be picked up by a parent, leaving little or no trace in the nest. Typically, a nestling will void one of these sacs after it eats, by wiggling its rear toward the edge of the nest. A parent watches this activity intently and is sometimes able to catch the sac before it hits the nest. The drawing at the opening of this chapter depicts this parental behavior. Before nestlings are feathered, the parent needs to stay on the nest to brood or shade them. So rather than leave the nest to carry the fecal sac away, the parent simply eats it. William Calder (1968) studied the urine concentration of the fecal sacs of roadrunners aged fourteen to twenty-six days. He found that the young roadrunner is not able to concentrate urine as well as the adult, for its kidneys are still developing. Thus a nestling fecal sac has more water than an adult dropping does. Calder estimated that, in a nest with five nestlings, each parent could get about 7.5 percent of its daily water requirement of four to six ounces by consuming nestling fecal sacs. He examined fecal sacs only from older nestlings, so it is probable that even more water could be in the sacs of younger nestlings. Parents I watched ate fecal sacs from nestlings less than ten days old but rarely ate them after most of the young were more than ten days old. When a parent no longer needs to brood the young, it usually carries the sac away rather than eating it. Parents take these sacs more than a hundred feet from the nest, probably to prevent a predator from finding the nest area. These sacs are about three-fourths-inch long, partially white, and easily seen. Finding a fresh fecal sac on the ground is a good clue that a nest is nearby. By placing white paper the size of a large fecal sac in the nest while the parent was away, I investigated how thoroughly the adult bird would clean its nest. I also put scraps of white paper on the ground near the nest. On its return, the parent immediately removed the paper from the nest and left the nest to carry away the paper on the ground. A parent would remove all white paper placed within about fifty feet of the nest. This compulsive cleaning occurred at the three nests I tested. The parent also carried away pieces of paper that were much larger than a fecal sac, including a large, white sandwich wrapper that a pedestrian dropped near a nest. The parent may have responded to the white color, rather than the size or shape of the object. Eltaher (1980) found that a roadrunner parent would remove a white Styrofoam egg from its nest, even though the parents accepted several nonwhite fake eggs. When the white egg was painted to match the off-white color of the roadrunner egg, the parent no longer rejected it. It may be that “white” signals “fecal sac” to the parent and elicits removal behavior.

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Courting While Caring for Nestlings Until the oldest nestling is twelve or thirteen days old, at least one parent is usually at the nest, brooding or shading the young, while the other is foraging for nestling food. Little courtship calling or display takes place during this time, but mating occurs occasionally when the male returns to the nest with food. His distant coos may let the female know his intentions. She may clack or bark in response. Despite the lizard or other nestling food in his bill, the male is able to produce a normal-sounding coo. During mating, the female still takes courtship food from the male but afterward returns to the nest and feeds it to one of the young. When nestlings are older, both parents spend long periods away from the nest, separately foraging for themselves and for nestling food. Sometimes they meet at the nest with food and take turns feeding nestlings. Once both parents returned with nestling food and mated before proceeding to the nest to feed the young. Plate 10 shows this mating, with the male holding a lizard, and the female holding a large green caterpillar. This increase in mating coincides with the pair’s beginning a second or a third nest. They may spend part of the time building a second nest, feeding young at their first nest, and feeding fledglings from the first nest. Other courtship behaviors that occur during this time are stick offers and whine calls, both associated with new nest building, and growl calls.

Caring for Fledglings Leading Chicks from the Nest Fledging is a process that happens gradually over several days. The average fledging age is nineteen days, but this age can vary with nesting circumstances. The youngest nestling of one clutch tarried in its nest until it was twenty-five days old. On the other hand, nestlings as young as seventeen days will fledge occasionally, especially if an older sibling is fledging at the same time. When the oldest nestling is about fifteen days old, the parents start giving a growl call that they will later use to lead young from the nest. The parent usually gives this call right before or after feeding a nestling. In the next day or so, the parent starts growling from the ground when approaching or leaving the nest. Often an older nestling will move out of the nest onto a nearby branch after the parent growls from the ground. Some nestlings return to the nest at night, while others perch on nearby branches until they fledge. The final phase of fledging occurs when a parent continuously growls from the ground beneath the nest until one or more nestlings climb down to the ground. Still growling, the parent moves rapidly away, leading the fledgling or fledglings to shelter in nearby shrubs, trees, or other protective cover. At a West Texas nest, a twelve-day-old nestling accidentally fell from the nest. The first day it perched in a nearby prickly pear cactus, but the parent moved it farther from the nest each day by using the growl call. The young bird clacked its bill continuously when the parent was away. The parents kept it regularly fed while they were feeding its siblings in the nest.

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It appears that the growl of the parent is the major and perhaps only stimulus that causes nestlings to fledge. When I played a tape recording of the growl near nests, older nestlings began moving about the nest or even climbed out of the nest to nearby branches, while younger nestlings gaped for food. At one nest, I was able to lead an older nestling to the ground by playing a tape of the growl call near the nest. When searching for fledglings in dense vegetation, I would find them by playing the growl tape. I would hear distinctive hollow bill clacking or see the fledgling moving in the brush in response to the sound. Once a wild fledgling ran from its hiding place and jumped on the tape recorder on the ground that was playing the growl call. Standing only a few feet away, I could scarcely believe that this young bird had ventured so close.

Sheltering and Feeding Fledglings For the first few days after fledging, young birds spend most of their time sitting on low branches or other protective cover, clacking continuously. They are rarely on the ground, except when a parent is nearby. A juvenile is very wary, ceases clacking, and remains motionless when a human is nearby. As a person moves closer to the bird, it freezes in an erect posture with its neck stretched out, bill pointed upward, tail pointed downward, feathers pressed close to the body, and eyes open (plate 17). In this position, it blends in well with surrounding branches. Young black-billed cuckoos assume a similar erect, frozen posture (Herrick 1910; Sutton 1940; Sealy 1985). Sealy noted that this behavior might allow a cuckoo parent to leave the rather helpless young for long periods to seek food for them over a large area. Parents regularly bring food to fledglings but feed them only about half as much as they do nestlings. Most of the food is now insects rather than reptiles. Several times during the day, a parent leads each fledgling to a new location by slowly walking away, while growling for the chick to follow it to a new hiding place. By the time they have been out of the nest a week, several fledglings usually stay together in or under shelter, waiting for food to come. After a few days of hiding, fledglings join a parent as it hunts for part of the day. They keep close to the parent, ready to grab food as soon as the parent catches it. At first the fledgling begs like a nestling—crouching with its head drawn into its shoulders, and gaping and buzzing (plate 18). The young start feeding themselves occasionally when they are about twenty-five days old, although most food is still supplied by the parents. When offering a large food item, a parent drops it on the ground in front of a fledgling, rather than placing the item in the young bird’s mouth. This behavior may help the fledgling learn to pick up and beat a large item on its own. Once I saw a parent drop a large lizard (apparently dead) in front of a young bird, pick it up, beat it a few times, then drop it again. The fledgling seemed to get the message, for it picked up and began beating the lizard. Parent and young took turns beating the food for several rounds before the fledgling finally swallowed it. Fledglings also gain other feeding skills during this time. They begin jumping into the air after butterflies, moths, and other flying insects. As they walk behind or along with a

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Figure 14. Thirty-five-day-old juvenile crouching after parental attack (parent in foreground).

parent, they continuously examine the ground, eating items. When about thirty days old, the young may hunt alone or with siblings for part of the day. By this time, a second nest may be occupying the parents for most of the day, so they have little time for the fledglings. At about this time, parents start vigorously attacking their young whenever they assume a begging posture. Sometimes a parent attacks its juvenile even if it is not begging. When attacked, a juvenile crouches, with head lowered and wobbling, and gives a low whine (fig. 14). The behavior and sound are similar to the whining crouch of the female during nest building. When about a month old, most fledglings are feeding on their own, primarily eating insects and other arthropods. They may occasionally forage with their parents but do not get much food from them. Adult foraging techniques, including wing-flashing to flush prey, appear in juveniles between thirty and thirty-five days old. If parents are nesting again, juveniles spend time around the new nest. This may account for some reports of multiple females at a nest. I never saw juveniles from a previous nest help with feeding or other parental duties at their parent’s new nest, but I cannot rule this out as a possibility. Juveniles of the groove-billed ani help their parents in their next nest. I wish I knew how long juveniles stay with their parents. Although adults attacked the older juveniles, the juveniles never left the territory of their parents and never stopped foraging with them completely. My latest field observations were of two-and-a-half-month-old juveniles that were still foraging with their parents for part of the day. It is likely that juveniles stay near their parents late into their first fall. Groups of three to six roadrunners seen foraging together in early winter could be juveniles with their parents. One biologist observed six roadrunners hunting together until late December (Gander 1958). There are no observations of such groupings in late winter or in early spring, so the family may disband for the winter. Roadrunners are able to breed their first spring (author’s observations on captive birds).

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Courting While Caring for Fledglings After all young are fledged, parents forage together again, unless they are incubating eggs or feeding nestlings at a new nest. The male’s courtship behavior increases dramatically, with coos, tail wagging, and presentations of food to the female, which may or may not respond. Perhaps the female decides whether renesting will occur, based on some environmental cue, such as food availability or a changing photoperiod. The pair give the growl often, but it is associated more with finding the fledglings than with courtship. Juvenile birds learn or refine future adult behavior through early experience. Juvenile roadrunners give some calls and displays of adult courtship soon after leaving the nest. These usually appear interspersed with playful chasing of each other, a common juvenile pastime. In this context, I observed wild fledglings cooing, growling, whining, tail-wagging, and offering sticks to one another when the birds were about thirty-five or forty-days old. It is likely that hearing their parents’ courtship calls and watching their visual displays encourage fledglings to call and tail-wag. Without the benefit of a parental example, my captive birds did not give any courtship calls until their first breeding season the next year. One exception barked as a juvenile. Captive juveniles did give visual displays similar to the tail wag, the flick, and the stick offer of the adults. These behaviors were isolated events, rather than part of the pattern of adult courtship.

7 Development of the Young

ven though starting as a half-ounce, helpless lump, a roadrunner will fledge when about nineteen days old and will be almost indistinguishable from its parent by the time it is two months old. The following chapter describes the major milestones in the life of a rapidly growing roadrunner chick.

E

Hatching I was elated when Niño’s parents, José and Pedra, produced an egg in the typical roadrunner nest they had constructed in their large outdoor enclosure in Norman, Oklahoma. Because they had not incubated any of their previous eggs, I placed this egg in an incubator. I was present when the egg hatched, so I can describe the details of this event. The first evidence of pipping was a slight crack near the larger end of the egg at 10:00 a.m. on the seventeenth day of incubation. By 8:00 p.m. that night, the crack was about one-sixthinch long. About an hour and forty-five minutes later, a tiny chunk of shell centered on this crack had been knocked off, revealing the inner membrane of the egg. No further pipping occurred until 6:30 a.m. the next day, when another crack started at the edge of the previous crack. Around 9:00 a.m. the inner membrane in the crack was open, giving a first glimpse of the tiny, black-skinned body inside, which appeared to be heaving regularly. This heaving could have helped the chick saw through the shell with its bill. The hatching process then moved quickly. By 9:07 a.m. the chick had extended the crack completely around the shell so that one-third of the shell separated from the remainder

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as the chick pushed and kicked its way out. As the shell separated, the chick’s head was visible at the cracked end, with the end piece of shell resting on its head like a beanie. Its small body was still heaving regularly. Within five minutes the chick was completely out of the shell—a coal black lump with scattered white hairlike strands covering its body. The entire process from making the first faint crack to breaking out of the shell took about twenty-three hours. After all the effort of hatching, I expected Niño to be exhausted and ready to rest, but immediately the hatchling began scooting about the incubator by pushing its feet and legs to one side and backward, with its tiny head and body supported by the floor of the incubator. Faint grunts and oinks sounded, as the chick made its first moves in its new world. It moved rapidly over a circular area about eight inches in diameter for several minutes, as if searching for something. If it had been in a nest in the wild, it could have easily fallen out of the nest. Its movements stopped whenever I lightly touched its back or covered it with a tissue. In the nest, a hatching chick would be confined beneath the incubating parent’s body. My touch may have simulated the presence of the parent’s body. Within the next hour, Niño gaped for food whenever I opened the incubator door. The bright reddish pink mouth, fluttering wings, and persistent buzzing noise told me that the bird was hungry.

Nestlings The newly hatched chick weighs about one-half ounce and is naked and black-skinned, except for pink skin on the rear mandibles and chin area. The bill, legs, and feet are also black. The white hairlike strands that stand out against the black skin of the hatchling are actually modified down feathers that mark the locations where feathers soon emerge. Birds do not have true hair like that of mammals. These strands cling to the tips of the bird’s feathers after it fledges, and their presence is a good way to tell a juvenile from an adult roadrunner. Plate 16 shows four wild nestlings, ranging from one to four days old. Another distinctive feature of the newly hatched chick is a tiny white bump (egg tooth) near the tip of its upper bill. The egg tooth, typical of a bird hatchling, is used to crack the shell during hatching. It stays on the young roadrunner’s bill until it is about twelve days old. The hatchling also has a bare black area behind the eye that becomes the blue, white, and bright orange patch of the adult. Plates 16 and 19 show the egg tooth and bare skin area in nestlings. The weight of a nestling’s legs and feet is about 25 percent of that of an adult’s feet and legs, although the nestling’s body weight is only about 5 percent of that of an adult. Plate 20 shows the large foot and the small wing of a four-day-old nestling. Despite its large feet, the nestling is not able to stand and walk until it is about eleven or twelve days old. However, on a flat surface it can scoot about by kicking and pushing with its curled-up toes, while resting on its belly and tarsi.

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The sightless, day-old nestling rests under the brooding parent most of the day, moving only to gape for food or to defecate. It instantly gapes in response to nest vibrations produced when a parent jumps on or off the nest. Although its eyes are open by the second day, it does not respond to visual cues until around the fourth day. Up to that age, it will gape when the nest shakes or when touched on its back, but it does not gape at moving objects. A nestling first follows movements by moving its head when it is between seven and eight days old. Tiny, white, backward-pointing papillae are scattered over the back of the tongue and the roof of the mouth. Two white areas on the rear of the upper mouth and on the rear of the tongue stand out when the mouth is wide open. Many altricial nestlings have a brightly colored mouth with these types of markings, called directive markings, which help the parent direct the food into the gullet of the nestling. The roadrunner nestling makes a buzzing call as it gapes and flutters its wings. This buzz is faint in the newly hatched bird but increases to a loud noise as the nestling gets older, sounding like the buzzing of bees. Around seven days of age, the nestling starts making a soft, bill-clacking sound similar to that of the adult, but with a hollow tone to it. Barely able to hold up its wobbling head, the day-old hatchling would seem to require only small pieces of food. However, it is able to swallow a small lizard or mouse right after hatching. Sometimes the lizard, snake, bird, or mouse may be almost as large as the nestling. The nestling swallows large items whole and headfirst but has to partially digest the swallowed end before it can gulp down more of the item. A nestling less than five days old takes from one to three minutes to swallow the main body of a lizard or bird, but often as much as thirty minutes is required to finally swallow the tail or tail feathers. By the time the nestling is nine or ten days old, it can swallow a large lizard, including the tail in less than three minutes. Taking my cue from the typical roadrunner parent, I fed small mice to Niño within several hours after it hatched. I had to hold the mouse to support it while Niño swallowed, until Niño was around nine days old. A nestling gradually gains control of its wobbling head, allowing it to orient its head toward the feeding parent. When about nine or ten days old, it assertively begs and grabs the food from the parent, even though it is able to move about only with its weight supported by its belly. At this stage the tail feathers are emerging, sheathed in a thin membrane that soon flakes off as the nestling starts to preen. From the half ounce at birth, the bird’s weight increases twelve-fold to around six ounces when it is fifteen days old. With the feathers out of their sheaths, the fifteen-day-old looks like a miniature adult for the first time, except for its relatively short tail, the growth of which lags behind that of the feet, legs, and bill. The white strands on the tips of the young bird’s feathers are still very visible and the formerly black bare skin around and behind the eye is tinged with blue and pink (plate 21). As fledging time approaches, the nestling sleeps less and spends more time moving about in the nest than when it was younger. It may move to a nearby branch for most of the

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day. When a parent is not present, older nestlings give a soft, hollow bill clack almost continuously. Hints of adult behaviors appear in older nestlings, although the nest constrains their movements. Sunbathing, prolonged preening, and sand-bathing motions are common at the late nestling stage. Now the young bird can walk supported only by its feet. My captives started to walk on their feet when about twelve days old.

Fledglings By twenty days of age the young bird weighs about nine ounces and is usually out of the nest. Although almost as large as an adult, it still depends on its parents for food. When the juvenile is about twenty-six days old, its back feathers are no longer tipped with the white strands, making it hard to distinguish the juvenile from an adult. At close range the strands may be seen still attached to the tip of the crest feathers. Plate 22 shows a twenty-three-day-old fledgling with the white strands still attached to the feathers of its back. Juvenal plumage is generally duller than that of an adult, having fewer distinct markings and lacking the metallic bronze gloss of adult plumage (J. M. Hughes 1996a). These characteristics may be seen with the bird in hand but are difficult to discern in the field, especially under varying lighting conditions. I found that the best way to identify a juvenile less than two months old is by the relatively short tail. The tail does not reach the adult length of ten inches until the bird is more than two months old. At close range, the pink mouth and the lack of an eye ring will confirm the bird’s age as less than two months. To see the pink mouth requires the patience to wait until the bird opens its mouth when panting or yawning. Another way to identify a juvenile is by its slender appearance and tentative behavior relative to those of an adult, but this means of identification requires field experience with the species. When a juvenile is around two months old, black specks and splotches in the pink mouth and the light buff eye ring of the adult begin to appear. The juvenile won’t sport the distinct eye ring of the adult until about seventy or seventy-five days old. Plate 23 shows a twomonth-old juvenile that looks like an adult but has only a faint eye ring. Also, a few white strands cling to this individual’s crest feathers, another clue that the bird is a juvenile. Plate 24 shows the distinct eye ring in a seventy-five-day-old juvenile that still has the mottled pink and black mouth and tongue of a juvenile. The mouth does not become entirely black until the bird is about eighty-five days old. After this time, it is especially hard to distinguish juveniles from adults. There are also subtle plumage differences between the juvenile and the adult before the juvenile’s first fall molt. These include the shapes and color patterns of the outer tail feathers and the outer primary coverts and can be detected only with the bird in hand (Pyle 1997). The juvenile roadrunner goes through its first feather molt during its first fall. My Oklahoma captives that hatched in June molted their first fall between late September and late November. All of their tail feathers and most of their wing feathers were replaced. They

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began molting again the following March. Adult roadrunners undergo an annual prolonged molt that begins in the late spring or early summer (J. M. Hughes 1996a).

Aging Young Roadrunners My captive Niño grew rapidly, reaching the weight of a small adult (ten ounces) at around one month old. Records kept by Kerry Muller (1971) for a nestling hatched at the National Zoo in Washington, D.C., showed a similarly rapid weight gain. Nestlings in the wild also showed rapid weight gains, as reported by Leon Folse (1974) for South Texas nestlings and Robert Ohmart (1973) for southern Arizona nestlings. Niño’s feet were almost adult size at one month old. The bird’s bill growth was not far behind, reaching adult size by forty to forty-five days of age. Tail feather development lagged, not reaching adult size until Niño was about sixty-five to seventy days old. Agility of the feet and the bill are critical to the success of a fledgling roadrunner because it must begin moving around and feeding itself soon after leaving the nest. The tail is also important for balance during running and fast maneuvering for food but may not be as essential to early survival as are the feet and the bill. To aid in future field research, I have included summary descriptions of behavior and measurements of various physical features of nestlings and fledglings in tables 3 and 4. These measurements are based on captive hatchlings that were fed abundantly and should be viewed as maximum values. When I raised Chico and Grande, my first captives, I was unaware of the tremendous amount of food they needed. Although these nestlings survived my inept mothering attempt, their growth was erratic and well below that of my later hatchling Niño and the National Zoo chick. Chico especially lagged behind, weighing at age twenty days what Niño weighed at eight days. That Chico eventually became an average-sized adult suggests that roadrunner nestlings can survive some food shortages in the wild and still reach normal adult weight. From my observations, weight is not a very good measure of nestling age. Bill, toe, or tail lengths are probably better indicators of age than is weight, but these measurements also can lag behind the averages if the bird is on a sparse diet. My measurements on toe, bill, and tail length of six captive young showed wide variability. These features grew much more slowly in the poorly fed Chico than in the well-fed Niño. This finding highlights the need to know the relative food supply at nests one is studying. Data on wild nestling weights and feather development at nests in Arizona were reported by Robert Ohmart (1973).

Temperature Control In the last chapter we learned that until the nestling is well feathered, parents need to brood it during cold times and to shade it during hot times to keep its body temperature stable. Temperature control in the nestling develops gradually, as its nervous system, hormonal system, and feathers develop.

Table 3 Development of nestling roadrunners from hatching to twenty days old Age

Physical appearance

1 day

Black skin, legs, and feet. Toes curled. No feathers or down; only stiff white strands (“hairs”). White egg tooth on tip of upper bill. Mouth bright reddish pink. Eyes closed to partially open. Bare black area behind eye. Tips of primary wing feathers and tail feathers starting to emerge. Eyes fully open and solid dark brownish black. Bill with egg tooth. Tips of secondary wing feathers starting to emerge.

2–3 days

4 days

5–6 days

Emerging feather sheaths on all feather tracts. Two middle tail feathers free from sheaths and about 0.5–0.6" long.

7–8 days

Tail feathers forming, about 1.0–1.6" long. Black iris appears. Black skin behind eye starting to change to dark reddish pink color. Feet gray. Middle tail about 1.6–2" long.

9–10 days

11–12 days

13–14 days

15–16 days

Feathers nearly all out of sheaths, with little bare body skin showing. Feathers totally sheathed only on upper neck, chin, and sides of crest. Bill 1.2" without egg tooth. Tail 2–2.5" long. Legs and feet gray. Tail 2.5–3" long. Bare area on side of head light pinkish orange; blue appearing around eye area. Tail 3–3.7" long. Looks like adult for first time (except for short tail).

17–18 days

Tail 3.7–4.5" long.

19–20 days

Tail 4.5–4.9" long.

Behaviors/physiological functions first observed Begs for food with gaping mouth, wing-flapping, and buzzing noise. Begs to vibrations only. Moves by pushing feet to one side and scooting on belly and tarsus. Toes curled. Gular flutter. Head wobbles. Nasal salt formation first seen.

Moves about more, still on belly and tarsus. More control of head wobbling. First attempts to reach feathers with bill jabs (like later feather-preening movements). Begins to beg to visual movement. Very alert. Begs vigorously, with wingflapping, gaping, and jumping. Strong toe-grasping response. Produces rasping, hissing sound. Head no longer wobbles. Can raise up on legs, still supported by belly. Eyes first follow hand movement. Pecks at tiny objects. Stands on feet in zygodactyl position. Still supported by belly and legs. Soft, hollow bill-clacking sound. Grabs food from parent or human. Moves about in nest. Stretches legs and wings. Crouches to birds flying overhead. Can walk supported only by feet. First fear of humans. Refuses food from humans. Makes louder hollow clack sound. Crest raises/lowers. Exposes bare skin behind eye. Moves about in nest. May leave nest to sit on nearby branch. Picks up food. Sand-bathing motions. Captives run on ground. Can run with full leg stride. Suns, exposing black patch on back. Picks up food. Able to preen feathers. Produces adult bill clack sound when startled Gives hollow clack almost constantly. Stops producing nestling fecal sac.

Sources: Muller 1971 (captive hatchling); Ohmart 1973 (wild nestlings in Arizona); author’s unpublished data (wild nestlings in West Texas and New Mexico and hand-reared nestlings from West Texas and southern Oklahoma). Note: Measurements should be viewed as maximums, given that some are from nestlings hatched in captivity with ample food supply. Weights not given, because they are highly variable with type and amount of food. The first nestlings hatched may develop faster for their age than the younger nestlings.

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Table 4 Development of fledgling roadrunners from twenty-one days old until first fall season Age

Physical appearance

Behavior first observed

21–25 days

Looks like small adult with short tail. Tail 4.7–5.3" long. Natal white strands attached to feather tips. Solid pink mouth. Solid dark eye; no eye ring.

Submissive crouching, head wiggling, and whining as begs from parent. Prostrate sunbathing. Stick-offer display.

26–30 days

Tail 5.4–5.7" long. White strands no longer on feather tips, except on crest feathers.

Seldom begs for food. Can catch insects. Beats larger food items.

31–40 days

Tail 5.7–8" long. White strands on tips of some crest feathers

First regurgitates food pellets. Tail wag. Flick display. Coo. Growl call. Wing-flashes.

60 days

Tail 9.8–10" long. Faint eye ring. Some white strands cling to crest feathers. Mouth mottled pink and black (plate 23).

Catches small vertebrates. Beats food and swallows whole.

75 days

Adult eye ring present (plate 24). Mouth mottled pink and black.

85 days

Looks like adult at distance. Mouth totally black. Adult eye ring present. Gray iris.

Bark call. Single coo call. Vertical tail flick.

First fall

First molt

Possibly still foraging with parents.

Source: Muller 1971; author’s observations (wild fledglings in West Texas and New Mexico and hand-reared birds from West Texas and southern Oklahoma). Note: See note for table 3. Refer to Pyle (1997) for differences between juvenal and adult plummage when the bird is more than 85 days old and before first molt.

The body temperature of an unfeathered nestling changes with the air temperature around it, making it more like a “cold-blooded” reptile than a “warm-blooded” bird. The unfeathered, black-skinned roadrunner nestling will quickly die of overexposure to extremes of cold or heat. Spending even a short time in direct sun can be fatal to an unfeathered nestling, for its black skin absorbs heat much faster than does light-colored skin. When shaded in above-normal heat, the unfeathered nestling can cool off by gular flutter, just as adult roadrunners do. I observed gular fluttering in my hatchling Niño when it was six hours old in a heated incubator. In a study of temperature control at a nest in Arizona, Robert Ohmart (1973) implanted a telemetry unit in a day-old nestling to measure its core body temperature. He found that the nestling’s temperature remained steadily at or near 104°F when the parent was brooding, but fluctuated when the parent was gone. Between three and four days old, the nestling was able to keep its temperature below air temperature during the hottest times of the day by losing heat through panting and gular fluttering. When air temperatures dropped below normal at night, the male’s brooding kept the nestling’s temperature stable.

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The nestlings that I raised did not have the luxury of an attentive, brooding parent to keep their body temperatures stable. Even the older, feathered nestlings (ages 9–14 days old) did not maintain a steady 104°F temperature in air temperatures lower than this. Their cloacal temperatures were as low as 73°F (nine days old) to 88°F degrees (fourteen days old). This suggests that up to at least fourteen days of age a roadrunner nestling cannot maintain a stable body temperature independent of the surrounding air. When temperatures are warmer than 104°F, young birds can keep their temperatures near normal by losing heat through panting and gular flutter.

Water Conservation In hot weather an adult roadrunner’s nasal salt glands help it avoid heat stress by conserving body fluids. This gland removes salt from the bird’s bloodstream and excretes it through its nostrils (see chapter 3). Robert Ohmart (1972) found that nestlings five days and younger had gland/body weight ratios that were from 2.5 to 3.9 times larger than those of adults. At this age, the unfeathered nestling’s bare black skin quickly absorbs heat, subjecting the bird to more of a heat load than an older, feathered sibling experiences. After a nestling is about nine days old, the salt gland develops more slowly, paralleling the development of feathers. I saw salt crusts in both captive and wild nestlings on their first day of life. Newly hatched Niño, raised in Oklahoma during a hot, humid summer, excreted drops of liquid from the nostrils. The salt crusts on the nostrils of a four-day-old chick are depicted in the drawing at the opening of this chapter. No studies have compared the effectiveness of the nestling salt gland with that of the adult gland, but the difference may be important to nestling survival in extremely hot weather. During hot summers in West Texas, nestlings had salt crusts on their nostrils more often than did their parents. The activity of the nestling’s salt gland is probably an important supplement to the work of its kidneys, which are not able to concentrate urine as well as an adult’s kidneys do.

8 Population and Range

n 1915, D. I. Shepardson predicted, “It is probably only a matter of time until the Road-runner will go the way of the Carolina Paroquet and the Passenger Pigeon unless public sentiment is aroused in his behalf” (1915, 160). This lament for the fate of the roadrunner appeared at a time when its population was subject to a serious human threat. Farmers and government wildlife managers were shooting roadrunners because of perceived threats to chickens, quail, and songbirds (McAtee 1931). Roadrunners were hunted as a game animal as well (Bryant 1916; Gorsuch 1932). Fortunately, the species did not become extinct as Shepardson predicted. Even as he wrote, the roadrunner was expanding its range eastward across Oklahoma, southern Kansas, and East Texas. Today this species is not listed as threatened or endangered anywhere in its U.S. range (see chapter 1).

I

Population Annual U.S. breeding bird surveys suggest that between 1966 and 2001 the overall roadrunner population may have slightly declined in the United States. However, these data have deficiencies because of the small sample size. Credible breeding bird survey data indicate that the roadrunner population has slightly increased in two subareas—the Chihuahuan Desert of western Texas and southern New Mexico, and the South Texas brushlands (Sauer, Hines, and Fallon 2003). The areas that consistently have the largest abundance of roadrunners in spring and summer breeding bird surveys and winter Christmas Bird Counts are in the southern parts

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Figure 15. Locations by county of roadrunner records from mid-1800s to present time.

of Arizona, California, New Mexico, and Texas (fig. 15). In these states, roadrunners are most common in the Sonoran Desert (Arizona), the Mojave Desert (western Arizona and Southern California), the Chihuahuan Desert (West Texas and southern New Mexico), and the South Texas brushlands (Sauer, Hines, and Fallow 2003). The only long-term study of populations is from the Rob and Bessie Welder Wildlife Refuge in the Gulf coastal bend region of South Texas. In this brush-grassland habitat, Leon Folse (1974) estimated a density of one breeding pair per 160 to 200 acres during his two-year study. His thorough mapping of roadrunner nests revealed that pairs were not uniformly distributed over a habitat but were clumped together in the more suitable habitats within the refuge—that is, in the shrubby, open areas. Other population density estimates come from counts of singing males or observations of pairs together, rather than from mapping nests. In a Sonoran Desert scrub habitat near Tucson, Arizona, one pair was estimated to occur for every fifty acres (Tomoff 1972). Only one breeding pair occurred on a forty-three-acre study plot in a mesquite bosque in the Mojave Desert (Austin 1970). Densities of singing males were from one per twenty acres to one per ninety-eight acres in shrubby grasslands of South Texas (R. R. Roth 1971).

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Roadrunners on the edges of urban areas may require only small breeding territories because of the abundance of food. I observed a roadrunner pair foraging over about twenty-one acres in a residential area in New Mexico. During this time, they raised two successful nests. Insects were plentiful on nearby vacant lots, which had native desert scrub vegetation. Adult house sparrows, bird nestlings, bird eggs, and pet food were major food sources for these parents. At another suburban nest I watched, the breeding territory covered about sixty acres.

Influences on Population and Range The rapid eastward range expansion of the roadrunner during the last century shows the ability of this opportunistic species to take advantage of favorable conditions when they appear. Although this movement could be explained away as simply a matter of time for the species to spread out from its original population center, the speed of the movement is remarkable. Also notable is that, coming from a desert habitat, this bird moved with apparent ease into humid, deciduous eastern woodlands. During the last hundred years, generations of roadrunners gradually moved across the central plains into the wooded habitats of eastern Kansas, eastern Oklahoma, and East Texas. From there, they continued eastward to the edge of the Mississippi River valley of Missouri, Arkansas, and Louisiana. This roughly five-hundred-mile movement is a remarkable distance for a ground-dwelling species whose individuals move only a few miles during their lifetime. I think that this unusually fast expansion of the roadrunner population into new eastern areas can be explained as a combination of three factors: the species’ long history as a dweller of open woods, rather than desert; the extreme flexibility of the species’ habitat, diet, and breeding habits; and human changes in the landscape that enhanced habitats for this species. The following sections discuss these factors as a background for understanding its range expansion.

Prehistoric Roadrunner Habitat Although in modern times the roadrunner has occurred mainly in desert areas, a look at its history as a species tells a different story. The roadrunner’s expansion into colder and wooded eastern areas may not be so remarkable when we consider that the bird’s history as a species is a great deal older than that of the southwestern deserts. Comparing the roadrunner with other western U.S. bird species whose lineages go back this far, the roadrunner is one of the few that now covers a larger area of the United States than it occupied in prehistoric times. The oldest-known fossil of the modern roadrunner, Geococcyx californianus, is from a cave in southeastern New Mexico and is estimated to be about 33,500 years old (Harris and Crews 1983). Other roadrunner fossils include twenty-five found in the Rancho La Brea tar pits in Los Angeles County (Howard 1962), several from the Southern California counties of Santa Barbara and Kern (Larson 1930), and several from northern Mexico. Paleontologists have

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determined that the bones of these fossil roadrunners are no different from those of the modern bird. However, we cannot assume that all aspects of the early roadrunner’s behavior and physiology were the same as the roadrunner of today. Fossils of the larger, extinct Conkling’s roadrunner are also reported from several southwestern locations and Mexico (see chapter 1). One of the most remarkable things about roadrunner prehistory is the types of habitat it formerly occupied. Plant fossils show that until about eight thousand years ago, open woodlands covered the areas that today are the desert scrub and desert grassland of the Southwest (Van Devender 1977). Woodland species included juniper, piñon pine, and evergreen oaks. Some shrubs of today’s deserts occurred in the woodland as well, including Joshua tree, desert almond, shad scale, and four-wing saltbush. The chuckwalla, the desert tortoise, and other reptiles that today are only found in desert habitats roamed these woods. The climate was cooler and the snow line was lower on western mountains than at present. Pine and spruce forest occurred at higher elevations above these woodlands, with tundra-like conditions on some of the highest mountaintops in New Mexico. Considering that for two-thirds of its history, the roadrunner was a species of cool, open woodlands rather than of desert scrub, a remarkable adaptation took place thousands of years ago, when its population survived the gradual development of desert scrub and grassland habitats that were emerging on the lowlands of the Southwest as the climate warmed. As a species, it never left its woodland home, however, for it still occurs and nests in open woods on southwestern mountains, especially in oak encinals, pine-oak woodlands, and piñon-juniper woodlands. Even in desert lowlands, roadrunners, like most other “desert” birds, are more common near wooded arroyos and riparian areas than in the midst of spindly, low desert scrub.

Roadrunner Adaptability In the above discussion, we saw how the roadrunner’s history characterizes it as much a bird of open woodlands as of deserts. But this information does not pin down the specific features of the bird that predispose it to a rapid invasion of new areas. To do this, we must glean clues from its everyday life. Just about every aspect of its natural history points to a predisposition to take advantage of new opportunities, as detailed in the following list: Omnivorous diet, eating everything dead or alive, plant or animal Typically long breeding season, with up to three clutches per year Nest site flexibility, allowing the use of any sturdy vegetation or artificial structures Complex courtship displays, including feeding of the female, which promote stability throughout the breeding season and may tie nesting to food supply Given this adaptability, the roadrunner population was ripe for expansion eastward as the landscape at the eastern edge of its range was opened up by human settlement.

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Humans and Roadrunner Range Expansion Although habitat destruction by humans is blamed for much animal extinction, the westward movement of pioneer farmers into the heartland of the United States ironically coincided with and probably facilitated the eastward expansion of the roadrunner through this area. The settlers’ modification of the landscape made better roadrunner habitats than had existed there before Euro-American settlement. The roadrunner’s needs are simple—open ground or short grass where it can feed, with tall vegetation nearby where it can roost and nest. If tall vegetation isn’t present, then about any aboveground object seems to work. The habitats in which roadrunners do not occur are characterized by one or more of the following features: Continuous tall grasses with no shrubs, trees, or sturdy cacti Closed woodlands or forests with a dense understory of brushy vegetation Desert scrub that is a monoculture of small, spindly shrubs such as creosote bush or sagebrush Large continuous wetland complexes, such as floodplain forests, wet meadows, and marshes These habitats do not provide the safety of tall vegetation for nesting and roosting, they lack open areas for foraging and nest location, or they are hard for a ground dweller to traverse. Some of the changes brought on by settlement that provided the open ground and tall vegetation needed for roadrunner habitat included: Clearing of the expanses of Great Plains grasslands and prairies in Kansas, Oklahoma, and Texas Planting trees in grassland areas around homes and along fencerows Clearing openings in closed-canopy woods in Arkansas, Louisiana, Oklahoma, and East Texas Providing roadway corridors through tall grass and woods Elimination of fires and bison, both of which had kept trees and shrubs at bay in the grasslands Draining wetlands for farming In addition, the roadrunner tolerates human presence very well, as evidenced by its survival in many suburban areas today (Webster 2000; Cornett 2001).

Range Expansion History Keeping in mind the factors discussed above, we now will look at the roadrunner’s trek across the south-central United States. Figure 16 shows the general areas of this range expansion.

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Figure 16. Range expansion of roadrunner from 1900 to 1975.

1830–1900 In 1835 the roadrunner’s known range was depicted as extending from the tip of Baja California, Mexico, to the San Francisco area (Ridgway 1916). At that time, most of the interior western United States had not been explored, so the roadrunner’s geographic range east of California was unknown. It was not until the 1850s that major U.S. government–sponsored expeditions provided a look at its range. These expeditions surveyed boundaries along the U.S.-Mexican border, explored potential railroad routes, and established wagon routes for pioneers moving westward. Most expeditions included army surgeons, who were often naturalists and compiled information on geology, paleontology, and biology. An interesting historical side note is that Spencer F. Baird, assistant secretary and later director of the Smithsonian Institution was the person who insisted that the surgeons selected for these expeditions also be naturalists. From these government reports (Baird 1859; Coues 1874; Yarrow, Henshaw, and Cope 1875; Bendire 1895; Ridgway 1916) and the records of the American Ornithologists’ Union (1886, 1910), it appears that the roadrunner’s known distribution prior to 1900 was in nine states: Arizona—statewide, but uncommon in the north California—statewide, except in high elevations and the northern tier of counties Colorado—southeastern, southwestern, and central areas Kansas—southeastern corner of the state Nevada— southern area of the state

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New Mexico—statewide, but uncommon in the north Oklahoma—southwestern corner of the state (Indian Territory) and in the panhandle Texas—in the southeastern, southwestern, and central areas and in the panhandle Utah—southern areas In 1853 John Newberry, an army surgeon-geologist for the Pacific Railroad Survey, provided a northernmost U.S. record for the roadrunner from Fort Reading, in Northern California at the upper end of the Sacramento Valley (Newberry 1857). This is still one of the northernmost U.S. records for the species. The northern edge of the roadrunner’s range extended from California through southern Nevada, southern Utah, central Colorado, and southwestern Kansas. The eastern edge of the bird’s range stretched from southwestern Kansas, western Oklahoma, and central Texas to the Gulf coast of Texas. The southeasternmost record was from Navarro County in north-central Texas in 1882.

1900–1940 Before the 1900s the eastern edge of the roadrunner’s range abutted grasslands and prairies from southwestern Kansas to South Texas. The demise of the buffalo and fire suppression had already allowed woody growth to invade these plains. Ranchers were the first settlers in much of the area, and the grasses were eventually overgrazed. A gradual shift from unfenced ranches to small farms took place as homesteaders came to plow the land. With settlers came roads and tree plantings for windbreaks. Overfarming eventually led to erosion and soil loss, resulting in the dust bowl days of the 1920s and 1930s. As farms were abandoned, more woody vegetation invaded the fields. The plains then provided an ideal habitat for the roadrunner—a mix of open land with shrubs or trees. In Oklahoma, changes came swiftly after the territory was opened for Euro-American settlement in 1889. Oklahoma ornithologist Margaret Morse Nice (1931, 22–23) wrote of the landscape changes that came with this settlement: “On April 19, 1889, the white man swarmed down on the land, much of whose wild life was still undisturbed. . . . Since then in eastern Oklahoma the primeval forests have been largely cut down, while in western Oklahoma the prairies have been turned into farms, towns, and cities.” She listed many grassland birds that quickly declined in number or became extinct in Oklahoma because of these rapid habitat changes. However, she noted that some insect-eating birds of open country benefited by the trees and shrubs in the former prairie areas. She reported the roadrunner to be present in all but the eastern third of the state in the 1920s. Another observer, Walter Colvin (1935), reported a roadrunner sighting in southeastern Oklahoma in 1920. In 1934 a nest was reported as far northeast as Arkansas City in south-central Kansas (Colvin 1935). The first Arkansas record was in 1936 from Hempstead County (southwestern part of the state). In Texas the blackland prairie, a narrow prairie band that extended from north-central Texas to south-central Texas, was plowed for cotton farming before the Civil War, and its rich soil was depleted before the Great Depression. Landowners converted much of this

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land to pasture, which was soon invaded by shrubs and trees. My mother grew up on an East Texas cotton farm just east of this prairie belt in the early 1900s and remembers that there were neither trees nor roadrunners on this land when she was young. Around 1930 the land was converted to pasture. Today oak groves dot these pastures, and roadrunners occur there. The first roadrunner record from this area was in 1936. By 1940 the roadrunner population had extended eastward to south-central Kansas, southwestern Arkansas, and northwestern Louisiana (James and Neal 1986; Lowery 1974). It had also moved northward in western and central Kansas to the Arkansas River.

1940–1980 During the 1940s roadrunners appeared in northwestern Arkansas and in central and southcentral parts of the state in the 1950s. In the 1960s this species advanced across the Ozarks to the Mississippi River basin in northeast Arkansas (James and Neal 1986). Roadrunners were first reported in southeastern and east-central Kansas in the 1950s (Brecheisen 1956). The first Missouri record was in 1956 near Branson in southwest Missouri (Brown 1963). In the late 1960s roadrunners first occurred in north-central Louisiana (Goertz and Mowbray 1971). The roadrunner advanced eastward across Louisiana, Arkansas, and southern Missouri until about 1975, when it reached its current easternmost boundaries near the Mississippi River basin. It also advanced northward to the Missouri River in central Missouri by 1975 but later retreated after three severe winters in the late 1970s (Norris and Elder 1982). The range of the Missouri population declined from thirty-six counties in the southern half of the state in 1976 to ten southern counties by 1978. Recently roadrunners have started moving northward again in this state (Robbins and Easterla 1992). Many parts of the region where Arkansas, Missouri, and Oklahoma meet are dominated by a cedar glade habitat. These glades consist of openings in oak-hickory forests where shortgrass prairie and eastern red cedar grow in rocky terrain that is strewn with boulders. Several other southwestern species reach their northeastern range limits in these glades, such as a collared lizard species (Probasco 1976).

1980 to the Present Time The only recent confirmed new roadrunner record along its northeastern range boundary is that of a bird that was seen near Lawrence, Kansas (Douglas County), in the 1998–99 winter (Kansas Bird Records Committee 2002). No further roadrunner sightings have been reported from this area. This record and other onetime roadrunner sightings that are out of the bird’s normal range probably represent single wanderers rather than a range expansion. Examples of “wanderer” records show up in figure 15, such as those in northern Colorado near the Wyoming border, in northwestern California along the Oregon border, in northern Utah, and in northern Kansas.

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Unconfirmed “out-of-range” reports of the roadrunner come from southeastern Iowa (Petersen 1970), western Mississippi in 1982 (Terence Schiefer, Mississippi Bird Records Committee, pers. comm.), southwestern Oregon in 1998 (Oregon Bird Records Committee 2003), and Manitoba, Canada (Koes 1991). The Canada sighting was undoubtedly an escaped captive bird.

Barriers to Further Expansion Given that the roadrunner’s range has not expanded significantly northward nor eastward since the mid-1970s, this roaming species may have reached its limits. It is probable that various combinations of factors have halted the bird’s range expansion. As these factors wax or wane over time, the local range can be expected to expand or shrink, as happened in the 1970s in central Missouri. On the other hand, with long-term climatic changes associated with global warming, the roadrunner’s northern range boundaries could gradually advance northward in the future. Taking into consideration the basic needs and survival mechanisms of the roadrunner, we can speculate on what environmental factors along the edge of its range may be at work to limit its advance northward or eastward. Keep in mind that a correlation of a factor with the edge of its range does not necessarily make for a cause-and-effect relationship. The following discussion of possible limiting factors raises as many questions as it answers, but I hope that it will stimulate further research into the roadrunner’s ecological relationships.

Rivers, Wetlands, and Bottomlands In Kansas the roadrunner’s northern range roughly coincides with the floodplain of the Arkansas River, although a few stragglers have reached points northward. In Missouri the northernmost records are to the south of the Missouri River (Norris and Elder 1982). Along the entire eastern range boundary of the roadrunner, an obvious physical barrier to any ground-dwelling bird is the vast Mississippi River basin, including the Mississippi delta in Louisiana. In east-central Arkansas and in Louisiana, the roadrunner’s range stops short of this river’s floodplains and wetlands. Roadrunners do not occur anywhere in the vast wetlands and bottomlands along Louisiana’s entire Gulf coastal area. Upland areas with open woods in southeastern Missouri appear to be good roadrunner habitat, but no roadrunner sightings have occurred there to date. A study of roadrunner ecology along the eastern and northeastern range boundaries would give us a better understanding of the species’ requirements and factors limiting expansion.

Climate In Atlas of Wintering North American Birds, biologist Terry Root (1988) examined the distribution and abundance patterns of North American wintering birds and the relationship of these patterns to environmental factors. She reported that the only factor that correlated

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with the roadrunner’s U.S. range was the number of sunny days. Its range roughly coincides with areas that receive at least 140 sunny days a year (clear days from sunrise to sunset). Other factors—average minimum January temperature, mean length of frost-free days, mean annual precipitation, average annual general humidity (pan evaporation), elevation, and structure of vegetation types—were not coincident with the roadrunner’s range. Root did find that the species is most abundant in areas of extremely low humidity (areas where the annual pan evaporation measures more than one hundred inches). The three factors that were most associated with other bird species’ winter range limits were average minimum January temperature, mean length of frost-free period, and vegetation. In addition Root suggested that the northern boundaries of wintering passerine birds (the roadrunner is a nonpasserine) appear to be limited by the energy demands of cold temperatures. The northern boundaries of passerine species seem to coincide with the point where raising their metabolic rate to about 2.5 times their basal rate would allow them to keep warm through the night. In the following discussion of climate as a limiting factor on the roadrunner’s northern range, I suggest that a combination of cloudy days, cold temperatures, prolonged snow cover, lack of woody vegetation, and scarce winter food is what limits the roadrunner’s northern range. I suspect that roadrunner populations and range boundaries are in a state of flux wherever the species encounters these conditions, some of which may vary from year to year. In the northern areas of the roadrunner’s range, cold temperatures and many consecutive days below freezing may lead to exposure deaths in roadrunners. Cold weather also causes food shortages and makes it difficult for birds to forage. Stomach contents from roadrunners in winter indicate that they feed on insects, insect egg cases, fruits, berries, and carrion (see chapter 3 for information on the roadrunner’s winter diet.) Several ornithologists have suggested that the roadrunner may enter a torpid state during extreme winter weather and food shortages. Although the roadrunner does become hypothermic on cold nights and possibly on cold, dark days, there is no evidence that it is torpid for long periods (see chapter 3). Thus it must continue feeding to survive. However, finding food in areas that have prolonged extremely cold temperatures and/or snow cover is another matter. Snow cover not only buries this food but also makes it difficult for a ground-dwelling bird to move about and forage. The only food items that roadrunners have been observed eating during periods of heavy, prolonged snow cover have been other birds, such as sparrows and starlings, and handouts from humans. Documenting the amount and length of snow cover at the northern edge of the roadrunner’s range from year to year would be interesting. In her atlas of wintering birds, Terry Root did not examine snow cover as a factor, because it is so variable and could be locally melted near a log or rock, making a feeding area available to most birds (Terry Root, Stanford University, pers. comm.). However, I think that finding isolated, melted patches in deep snow would be more of a challenge for the roadrunner than it would be for flying birds. During times of snow, I suspect that roadrunners in wooded areas forage primarily in

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bushes or trees and are more successful in finding food than roadrunners in open areas with only ground vegetation. Besides being a roadblock in the Midwest, cold temperatures and snow cover may hinder the roadrunner’s northern advance in the western states of Colorado, Nevada, Utah, and California. Cold temperatures, snow, and lack of woody vegetation in the four-corners area of Arizona, Utah, Colorado, and New Mexico may explain the low populations or absence of the roadrunner in these areas. Some areas where roadrunners occur have cold winters, snow cover, and scarce winter food, but these areas also have winter sunshine (at least the 140 sunny days from sunrise to sunset reported by Root 1988). Morning sunning behavior is used by the roadrunner to warm up from its energy-saving nightly temperature drop. On the other hand, midwestern winters are characterized by dark, overcast days, sometimes going for weeks without sun. This lack of sunshine could prevent the roadrunner from becoming warm enough to forage, thereby tipping the energy balance to quickly deplete fat reserves. The results of a lab study of daily activity patterns of roadrunners suggests another negative feature of cloudy, winter days (Kavanau and Ramos 1970). These roadrunners were less active on overcast days than on sunny days during the June and July study. Dim light and darkness inhibited movement of the birds, and bright light stimulated them. Kavanau and Ramos speculated that the roadrunner’s visual system is not suited to dim light.

High Elevations The roadrunner usually occurs below elevations of about seven thousand feet. High mountain ranges border the roadrunner’s range boundaries in the western United States, including the Rocky Mountains in Colorado and northern New Mexico and the Cascades, the Klamath Mountains, and the Sierra Nevada in California and Oregon. The Sierra Madre Oriental, the Sierra Madre Occidental, and the Sierra Madre del Sur also appear to be barriers to roadrunners in Mexico. Most likely, the same climatic factors of low temperatures, snow, and food scarcity, discussed previously, work against roadrunner populations in high elevations. On the other hand, roadrunners have occasionally been sighted at high elevations in the mountains of Colorado and New Mexico (Sutton 1940; A. M. Bailey and Niedrach 1965). They may forage in these higher areas for several months, as one did at more than eleven thousand feet during the 1956–57 winter in Colorado.

Lack of Nesting and Roosting Habitat As detailed earlier in this chapter, roadrunners do not occur in areas dominated by spindly, low shrubs, for the birds need a mix of open area with tall, sturdy vegetation for safe nesting and roosting sites. This habitat requirement may partially explain the species’ failure to advance very far into the Great Basin Desert areas of Utah and Nevada, which have primarily sparse, low shrubs and cold winters.

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On the other hand, this species does not occur in areas where the near-to-ground vegetation is too thick, such as in impenetrable woody thickets. Ornithologist Joe T. Marshall, Jr., found roadrunners in pine woodlands in Sonora, Mexico, but not in this same habitat in adjacent areas of Arizona. He attributed the difference to the open, parklike nature of the Mexican woodlands relative to the thick, woody undergrowth in the Arizona woodlands. In the Arizona areas, fire suppression had allowed the dense undergrowth to persist (J. T. Marshall 1957, 1963).

Urbanization Although roadrunners often thrive in suburban areas (Webster 2000; Cornett 2001; author’s observations), the roadrunner population declines as development overtakes natural features when extensive urbanization occurs. This loss of the roadrunner presence has occurred in California for the San Francisco Bay area, coastal Marin County, Santa Barbara County, and San Diego County (J. M. Hughes 1996a).

Parallel Range Expansion of the Armadillo A discussion on roadrunner range expansion isn’t complete without mentioning the range expansion of the nine-banded armadillo, which moved northward and eastward from the Texas-Mexico border in parallel with the roadrunner movement. The armadillo’s expansion coincided with the coming of Euro-American settlers to Texas. Grassland clearing, row-crop farming, prairie fire suppression, decreased hunting, and an invasion of shrubs and trees into former open grasslands are some of the factors thought to favor the armadillo’s expansion. Prior to about 1850 the armadillo was not found north of the Rio Grande in Texas. In 1905, biologist Vernon Bailey reported that it occurred in Texas eastward as far as the Colorado River in central Texas (Davis and Schmidly 1997). By 1914 it was in East Texas (Trinity River) and along the coast to the Louisiana line. By the 1970s the species occurred in Louisiana, Arkansas, and Oklahoma, and more recently has ventured into Kansas and Missouri. Another population, introduced in Florida, has spread westward to the Mississippi River. Unlike the roadrunner, the armadillo is not adapted to an arid climate and thus has spread no farther west than the eastern parts of the panhandles of Texas and Oklahoma. However, its northern range limits in Kansas and Missouri and its eastern limits at the Mississippi River so closely correlate with the range of the roadrunner that we may question if similar factors affect their ranges. The armadillo’s mainly insectivorous diet, supplemented by very small vertebrates, plant matter, and carrion is reminiscent of that of the roadrunner. A study of armadillo food habits that identified the stomach contents of more than eight hundred armadillos found that 93 percent (by volume) of their food was animal matter, mainly insects and other invertebrates (beetles, termites, ants, caterpillars, earthworms, millipedes, centipedes, and crayfish).

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Reptiles, amphibians, and birds’ eggs were present, but in very small amounts. Other occasional food included berries, fruits, fungi, and carrion (Davis and Schmidly 1997). Because the armadillo does not hibernate, is insectivorous, needs to eat daily, has very little body fat, and is not good at conserving heat, cold winter weather and snow cover may limit the armadillo’s northern range as much as they limit the northern range of the roadrunner.

9 A Bird of the People

y interest in folklore about the roadrunner began when I came across the following words of J. Frank Dobie in his classic article “The Roadrunner in Fact and Folk-lore”: “Perhaps no other bird of North America, excepting the eagle and the turkey, which the Aztecs had domesticated long before Columbus sailed, has been so closely associated with the native races of this continent” (1939, 4). Descriptions of roadrunner cures, superstitions, and folktales compiled from his years of studying southwestern folklore fill this fascinating article. It is not surprising that Native cultures existing side by side with the roadrunner for many centuries have incorporated the bird into many of their folkways and rituals. The roadrunner also was a favorite animal of the early Euro-American pioneers who settled in the Southwest. Western writer Arch Napier (1969), in an article entitled “The Clown That Met the Wagon Trains,” described how the roadrunner’s antics helped lift the spirits of these apprehensive newcomers to the desert. In her 1922 book on the birds of New Mexico, ornithologist Florence Merriam Bailey told how roadrunners would dart onto roads and race horses, keeping ahead of their trotting. The roadrunner remains a southwestern icon today, continuing this age-old association with humans. After exploring roadrunner folklore, I came to agree with Dobie that “any animal is interesting to man not only for the facts about him but for what human beings associated with the animal have taken to be the facts” (1939, 19). So I include this chapter on roadrunner folklore to complement the chapters of roadrunner facts.

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Pueblo Indian Folklore Many American Indian tribes of the Southwest knew the roadrunner and incorporated it into their culture. Folklore about this bird is best documented for the Pueblo Indians of northeastern Arizona (Hopi), of northwestern New Mexico (Zuni), and along the Rio Grande drainage in north-central New Mexico (nineteen pueblos, such as Acoma, Cochiti, Nambé, San Juan, Taos, and Sia [Zia]). The Pueblo people have lived in this region for many centuries. Clustered in small villages (pueblos), their society is based on communal farming and worship (Trimble 1993). Before the Spanish invasion, more than one hundred villages were scattered throughout New Mexico. In the Pueblo culture the powers of the roadrunner center around its courage, strength, and endurance, as well as its X-shaped track, thought to have power to confuse the enemy. Anthropologist Hamilton Tyler described specific historic uses of birds by Pueblo Indians in his 1979 book, Pueblo Birds and Myths. Grouping birds by their roles in religion, he placed the roadrunner in the “birds of war” group, along with nuthatches, wrens, woodpeckers, and jays. In the Pueblo context, conflicts with witches and ghosts of the dead are also considered war. Anthropologist and Zuni Ed Ladd documented the uses of birds specifically by the Zuni Pueblo in his 1963 master’s thesis entitled “Zuni Ethno-ornithology.” These two references are the main sources for the following discussion of Pueblo Indians.

Roadrunner Bravery The roadrunner’s bouts with rattlesnakes give the bird a reputation for bravery, explaining its use in ceremonies devoted to strength and knowledge. The rites of the Little Fire curing society at Zuni involve two persons representing the roadrunner, who hop and skip to the altar with an eagle’s wing feather in each hand. The Zuni curing society that treats rheumatic troubles and convulsions is always directed by a member of the Roadrunner Clan. Roadrunner feathers were used for courage in the scalp ceremony of the Zuni Pueblo (Parsons 1924). The scalp-kickers wore crossed roadrunner feathers in their moccasins and prayer feathers of the roadrunner in their hair. Anthropologist M. C. Stevenson described these scalp-kickers firsthand in 1904 (583–84): They wear their ordinary dress, with white blankets bordered in red and blue over their shoulders. The quill ends of two feathers of the chaparral cock—one an upper tail feather and the other an under tail feather—have been crossed and placed in line by their brothers in consanguinity between the second and middle toes of the left foot, the tips of the plumes pointing toward the foot, and the moccasins carefully drawn over. “The feathers give courage, for knowledge and courage come from this bird, who is the keeper of courage.” After the girls reach the scene, the same brothers tie similar but somewhat larger feathers to the left side of the head with a strand of the hair and a cotton string already attached to the plumes. The plumes must not be removed for four days.

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Roadrunner Swiftness Roadrunner swiftness is celebrated by the Pueblo Indians. Near the Sia Pueblo in New Mexico, the people run races around a mountain named after the roadrunner. At Hopi in northeastern Arizona, roadrunner feathers are sometimes tied to a horse’s tail for speed and endurance. Young Hopi men occasionally eat roadrunners to gain swiftness and endurance. Eating roadrunners to enhance running skills is also reported from the Tarahumara Indians of the state of Chihuahua, Mexico, famous for their long-distance running ability (Dobie 1939).

Roadrunner as Bringer of Rain Roadrunners are associated with prayers for rain in several Pueblo groups. The roadrunner kachina of the Hopi, called Hospoa, appears in only a few Hopi ceremonies, where it is associated with soliciting rain. One ceremony that always includes Hospoa, along with many other birds, is the Powamu, a bean dance or bean-planting ceremony (B. Wright 1977). Ethnologist Jesse Walter Fewkes described the roadrunner kachina in 1903 (98): Hospoa, the Road Runner . . . has a green helmet covered with rows of black and white crescents, a short beak, and stellate eyes. On the back this bird has a painted skin stretched over a framework, called a moisture tablet. To each upper corner are attached two feathers, which project horizontally, and along the edges is a string with attached horsehair stained red. There is a flute in one hand, a rattle in the other. The garments are a ceremonial kilt, girdle, and embroidered sash. The Zunis of New Mexico use feathers from seventy-two bird species on various types of prayer sticks (Ladd 1963). Specific feathers from different parts of a bird’s body may have different meanings. Each member of the tribe offers many prayer sticks during the year. Feathers from some species are used only on the prayer sticks of the spiritual leaders. These include roadrunner feathers, which are reserved only for the prayer sticks of the rain priests (Ladd 1963). It is noteworthy that in many cultures worldwide, various cuckoo species are associated with predicting or bringing rain and are called “rain bird” or “rain crow” (Ingersoll 1923). In the southern United States, yellow-billed and black-billed cuckoos are often referred to as rain crows.

Roadrunner Tracks for Protection Pueblos value the roadrunner’s X-shaped track because it appears to be heading in opposite directions at once (fig. 17). A Hopi man told anthropologist Polly Schaafsma (1989) that the roadrunner is a liar, because of its tracks. Drawings of roadrunner tracks are used to ward off spirits of the dead by confusing them as they look for the way to the village. Tracks also protect people from the dead, who

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Figure 17. Roadrunner track of right foot, pointing forward and facing to the right.

return on All Souls’ Day. Anthropologist Elsie Clews Parsons (1939, 859) gave the following description on All Souls’ Day at Nambé: “Before leaving the house each had taken a piece of charcoal and outside had marked on his left sole and on his left palm the cross-like track of the chaparral cock, so that the dead would not make him sick or do him any harm. The chaparral cock has power (piñan).” Roadrunner tracks also protect the dead from evil spirits and witches. At the pueblo of Cochiti in New Mexico, relatives scratched the roadrunner track’s X shape into the floor to encircle the body of the deceased (Dumarest 1919), providing “a magic circle for the purpose of preventing evil spirits or Brujos from finding out where the soul of the deceased goes,” thus protecting “that soul from their persecutions” (from unpublished 1888 journal of anthropologist Adolph Bandelier, quoted by Lange 1959, 416). Anthropologists Elsie Clews Parsons (1929) and Ruth Benedict (1935) described a ceremony at San Juan Pueblo in which roadrunner tracks were used to protect the altar during the installation ceremony of the winter chief. The tracks were drawn in the meal “road” (a trail of cornmeal). Roadrunner tracks were also painted on the altar wall at the winter chief installation ceremony at Tesuque Pueblo (Parsons 1929). In several of the New Mexico Pueblo groups, crossed pieces of yucca were laid as stepping-marks on an “altar” road to represent the X shape of roadrunner tracks. Their purpose was to confuse any witch that was pursuing a person (Parsons 1939).

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Other Symbolic Uses in Pueblo Culture The personal fetish represents its owner and is made by a society father and given to a new member on initiation into the society. Roadrunner feathers are used in personal fetishes by the Pueblo Indians at Acoma and Sia (Zia) in New Mexico. Many Pueblo rituals and ceremonies use birds to represent six directions: north, south, east, west, zenith (above), and nadir (below). Roadrunner feathers represent the nadir in Hopi and Sia ceremonies (Tyler 1979).

Prehistoric Pueblo Indian Culture Roadrunners have been part of the Pueblo religious expression for more than eight hundred years. Modern-day Pueblo peoples are considered to be descendants of prehistoric Pueblo people (Anasazi, or ancient ones), who were once widespread in the Southwest. The roadrunner and other birds appear to have been at least as important, if not more so, in Anasazi religious expression as they are in the culture of their Pueblo descendants. Although the Anasazi did not have a written language, they left many clues about their culture in their murals and prolific rock art, which give us a glimpse into their ceremonial uses of roadrunners. Pottery Mound was a Pueblo community in central New Mexico that thrived from about a.d. 1300 to 1475. Mural frescoes from the walls of seventeen excavated kivas (underground ceremonial rooms) are outstanding remains of prehistoric Anasazi art. The artifacts indicate Pottery Mound was a ceremonial center and a significant sacred site. Major features include a Mexican-style pyramid and many prehistoric murals. Bird feathers are the most common single motif in these murals. Feathers of seventeen bird species have been identified. Of these, the roadrunner was the fourth most common species represented, after the bald eagle, the raven, and the yellow-headed blackbird (Hibben 1975). The Anasazi also left roadrunner characters in their prolific rock art. Another nearby culture—the Mogollon peoples in southern New Mexico and adjacent areas of Texas—used the roadrunner track as a major symbol in its rock art. A few of these sites also have petroglyphs showing the entire roadrunner. Anthropologist and artist Polly Schaafsma (1989) investigated the use of roadrunner tracks in the rock art and ritual of this prehistoric period. In her interesting article “Supper or Symbol: Roadrunner Tracks in Southwest Art and Ritual,” she rejected previous interpretations of these tracks as merely being symbolic for obtaining food. She presents evidence that the roadrunner and/or its tracks had significant religious symbolism in these prehistoric southwestern cultures. Schaafsma found that roadrunner tracks were associated most often with carnivore tracks, especially those of the mountain lion. Besides appearing in rock art, the tracks of these two creatures are often together on other artifacts from the period. Figure 18 shows one of these petroglyphs from the San Diego Mountains of southern New Mexico. Gleaning

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Figure 18. American Indian petroglyph of mountain lion track and roadrunner track near Hatch, New Mexico. Photo by Curtis Schaafsma, provided by Polly Schaafsma.

clues from the modern Pueblo symbolic uses of roadrunners, Schaafsma concluded that the rock art tracks probably were important symbols of the bird’s courage and strength. The mountain lion often acts as a guardian and protector for war, hunting, and medicine societies. The occurrence of roadrunner tracks with mountain lion tracks may be an indication that roadrunner tracks also had guardian and protective functions. Roadrunner bones have been reported from prehistoric Indian ruins at Snaketown (near Phoenix, Arizona), dating from as early as a.d. 100. They appear at a number of later sites, including Grasshopper Pueblo, Wupatki, Tonto Cliff Dwellings, and the University Indian Ruin in Arizona; Gran Quivira, Pueblo Largo, Gila Cliff Dwellings, and Las Madres Pueblo in New Mexico; and Casas Grande in Chihuahua, Mexico (Longacre, Holbrook, and Graves 1982). Archaeologists are not sure of the significance of roadrunner bones at these sites, other than as a source of feathers for rituals and ceremonies.

Other American Indian Folklore Stories about the roadrunner are preserved in the oral traditions of other southwestern tribes, with some appearing in written form. An Apache story tells why the roadrunner is the leader

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of the birds. After eliminating several other birds for various reasons, the birds chose the roadrunner because it would be fast in running to meetings and it had good talking ability. The Pima Indians have a story telling how the orange color on the bare skin behind the bird’s eye developed (Curtin 1949). In one version the roadrunner flies to the sun, a four-day trip, to obtain fire for an old woman who wanted to cremate her pet rattlesnake. Lightning struck the head of the roadrunner on the return trip, producing the orange color. In the other version, Lightning Man wounds each side of the roadrunner’s head with gunshots.

Mexican Folklore I was able to explore Mexican folklore firsthand in the state of Chihuahua, Mexico, in the late 1970s. At that time folklorists considered the whole area of northeastern Chihuahua (south of Presidio, Texas, in the Big Bend area of Texas) to be an excellent place to investigate Mexican folk customs. Because of its isolation and lack of outside influences, its residents retained more of the old folklore than did more modern areas of Mexico. Likewise, this area of the Texas border is remote from large urban areas, having only the small towns of Presidio and Redford. At the suggestion of folklorist Joe Graham at the Institute of Texan Cultures in San Antonio, Texas, I visited the Texas border area with historian Enrique Madrid, Jr., from Redford as my guide and interpreter. Besides interviewing three curanderos (male folk healers) and one curandera (female folk healer), we also talked with five elderly Mexican men and women in the town of Ojinaga, Chihuahua, Mexico. Later I interviewed several elderly residents of another small Chihuahuan village, Santa Elena. These people appeared to have firsthand knowledge of the roadrunner in the wild, for they gave accurate answers to my questions on the natural history of the bird. When I played tapes of several common roadrunner calls, they were often able to tell me the context of each call. The following translation of a story written for me by an elderly Mexican man typifies the insightful observation of a person who has lived alongside the roadrunner. The Paisano It’s from the country. It eats lizards. It runs along roads. They look for food to keep their offspring fed. The paisano never rests. It spends all its time running along roads. It never rests all day long. El paisano is very healing. It is good for many illnesses. The meat of the paisano is very good and also the lard for sickness. I was sick and the healer prescribed this type of animal for me and I ground the paisano meat up very fine. This animal is very good for healing because it eats snakes and also seeds of wild herbs. The paisano likes very much to make its little house atop the mesquite so no one can bother its young. Here this story of the paisano ends. Written by Jacinto Hernandes Garsia, Santa Elena, Chihuahua, Mexico In the following paragraphs I summarize some of the folk-healing uses of the roadrunner. Folk sayings and tales I heard along the way are also included.

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A Potent Remedy The most common healing use described by the curanderos and laypeople alike was for healing skin problems, such as sores, boils, itching, hives, rashes, and stings. Eating a stew or soup made of whole roadrunner boiled with onions, garlic, tomato, cumin, and cilantro was the common prescription. Eating the roasted meat and rubbing lard of the bird on the skin were alternative treatments. One of the men claimed that eating three nestling roadrunners once cured his boils. Tuberculosis and other lung problems were thought to be cured with roadrunner stew. Two of the curanderos were firm about this cure, although their recipes differed. One insisted that rice must also be an ingredient. Another curandero used fresh roadrunner blood rubbed on the chest for lung problems. A roasted roadrunner was used as a blood tonic by the curanderos. A variation was to dry the roasted meat and grind it into a powder for sprinkling on food. One woman from Palomas, Mexico, stressed that roasted runner was especially good for anemic children with no appetite. In northern Mexico in the 1920s, West Texas photographer W. D. Smithers (n.d., 38) witnessed the roasting of a roadrunner by a curandero for use as a blood tonic: The bird was prepared by killing it without losing any of its blood, then it was taken to the river bank and completely covered with wet clay all over the feathers. In a hole dug in the center of a large fire the clay-covered bird was put in, covered with hot ashes and fire, to cook for about one hour. When removed from the fire, the baked clay was taken off in pieces and the feathers came off with the clay. The meat looked very appetizing, but the thought of it being cooked without being dressed spoiled the desire to try it. The entire bird was the prescription for the cure. Several laypeople used roadrunner lard as an ointment for earaches, but none of the curanderos used it. One man described how he had rubbed the lard in his hair as a cure for lice but ended up with even more lice. To cure a mental illness called susto, a tea is made from boiling roadrunner tail feathers. The feathers also make good throat swabs, according to one healer. When asked why the roadrunner is good for healing, all the healers told me that its ability to eat and digest venomous snakes, scorpions, centipedes, tarantulas, and other dangerous creatures is the source of its healing power. The laypeople likewise attributed the roadrunner’s healing power to its ability to eat venomous animals. One of the curanderos had two additional reasons for its power—its ability to live a long time without water and a unique digestive organ that it has. Because the roadrunner fights rattlesnakes and survives, it is thought to know a remedy for snakebite. Several similar tales relate how a roadrunner will pluck and eat the leaves of a certain plant after being bitten by a rattlesnake. Unfortunately, my translator was not familiar with the Spanish name for the plant, and it couldn’t be identified. The curanderos all stressed that the roadrunner is a potent cure and that only one bird is needed, which is fortunate for the roadrunner population. Since none of these healers had

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roadrunners or roadrunner parts on hand to show us and had other remedies for these same ailments, I asked how common its use really was. The curanderos said they did not catch a bird until they needed it. A young Ojinaga pharmacist expressed a skeptical view of these roadrunner folk cures. He commented that those who can catch a roadrunner don’t need a cure.

Roadrunners as Pets and Food Years ago roadrunners were occasionally kept as pets in Mexican homes. Like a barnyard cat, pet roadrunners would keep homes free from mice, insects, scorpions, and other undesirable pests. They also were kept on hand for healing, as the need arose. In a 1865 article on the birds of South Texas, ornithologist H. E. Dresser wrote: “The Mexicans often keep this bird in a semi-domesticated state, in order to kill them in case of sickness: for they firmly believe that their flesh is a certain cure for many disorders” (1865, 467). People whom I interviewed on both sides of the Rio Grande border said that roadrunners are not kept as pets today. Two Mexican people said that they ate roadrunners for food, but only rarely. One said that even though it doesn’t look appetizing, it is very tasty. John Newberry, a surgeon-geologist with a U.S. government railroad survey, reported in 1857 that the roadrunner was a food item in California: “It is frequently brought into the San Francisco market and is reported very good eating” (1857, 91).

Sayings and Superstitions A common saying along the border referred to the bill-clacking sound of the roadrunner. I heard several versions of this saying from people on both sides of the border: “No hay para que tonar el pico,” which meant “There is no reason to get excited.” Literally, this version translates as “There is no reason to snap your bill.” Another version is “Ya basta, vete a tonar el pico,” which translates as “Enough, go clack your bill somewhere else.” The latter is used to get rid of bothersome children. A short version is “No trina el pico,” or “Don’t get so excited”; the literal translation is “Don’t rattle your bill.” Fittingly, the most common superstitions I heard were tied to roads or trails. If a roadrunner crosses in front of you on the road, you will have a bad trip or bad luck in general; if the bird turns back when partway across, you will have a good trip or good luck. If you are following a roadrunner on a road or a trail and it leaves the path, bad luck will follow. J. F. Dobie (1939) reported a different version of this superstition from Indians in Mexico: the roadrunner brings good luck if it crosses the road from left to right, but bad luck if from right to left. One of the elderly Mexican men I interviewed had spent years as a goatherd and a soldier in very remote parts of Michoacán in southwestern Mexico. This ninety-year-old man told me that because the roadrunner travels all the paths, it will lead you to a lost person. His parents told him a story about a roadrunner leading them to a place in the desert where the

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body of a missing man was found. J. Frank Dobie (1939) reported another version of this superstition from Mexico: if you are lost, find a roadrunner and follow it, and it will lead you to a trail. One of the curanderos used some of the roadrunner’s behavior for weather forecasting: when a roadrunner is seen stocking up on food in the fall, a bad winter is coming; when the bird calls from a high perch early in the morning, it will rain that day. This reminded me that the clucking call of the yellow-billed cuckoo is also thought to forecast rain, and one of its folk names is “rain crow.” (Note: There is no field evidence of roadrunners storing food.)

Stories about the Roadrunner Most of the folk stories about the bird probably started with an observation of it doing something unusual that became greatly embellished over the years, for there is a grain of truth in many of the stories. In times past, these tales were widespread throughout the Southwest and Mexico, not only among Mexicans and Mexican Americans but also among the EuroAmericans who lived alongside them. We will look at a few of these stories.

A Peculiar Hybrid With a mischievous twinkle in her eye, an elderly Hispanic woman from a ranch near Redford told me: “Roadrunners are sort of rascals because they do mess around with chickens.” That male roadrunners mate with domestic hens appeared to be common knowledge among the people I met along the border. Most described the offspring as a small chicken with a large crest. One man said that the chicks were more paisano (roadrunner) than gallina (hen). J. F. Dobie (1939) reported tales of a similar cross between a roadrunner and a game chicken to produce a superior fighting cock. This folktale probably started because roadrunners often mix with chickens in their pens. Roadrunners have even been reported roosting in sheds with chickens. However, roadrunners are there not to court the chickens but to feed on the many insects that thrive in chicken pens (Jensen 1923).

Reverent Roadrunner Ornithologist Harry Oberholser (1974) related that some Mexican people in Texas believed the roadrunner to be a Christian because of the X shape of its tracks, which resemble a cross. Others believed it to be a Christian because it bows its head to pray every day (Dobie 1939). The head-bowing posture of the male roadrunner during its coo (see fig. 7) or tail-wag display (see fig. 9b) could have started this tale, or possibly one of the bird’s sunning positions, which it holds for many minutes as if in a trance, is the origin of the tale (plate 8).

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Roadrunner Rancher Many lizards escape from predators by losing their tail in the jaws or bills of their captors, but the lizards soon grow new tails. A popular story depicts roadrunners keeping herds of lizards for food. Instead of eating the whole lizard, the roadrunner eats just the tail, ensuring a steady supply of tails for itself and its family.

Corral of Thorns The story of the corral of thorns is probably the most often told and most often debated roadrunner folktale of all time. This tale is found in Mexican, Hispanic, American Indian, and Western folklore, both in the United States and in Mexico (Lewis 1897; Dobie 1939, 1978). The basic story is of a roadrunner that comes upon a sleeping rattlesnake. The roadrunner builds a corral of cactus joints around the still-sleeping snake. The bird then begins pecking at the snake to waken it. To escape the roadrunner’s attack, the snake frantically thrashes about, striking itself on the cactus spines. The snake eventually dies either from being pierced by the spines, from blows of the roadrunner’s bill, or from biting itself in frustration. One version has the roadrunner using a cactus pad like a bullfighter’s cape to lure the snake into piercing itself on the spines. From articles I read about this tale in several bird journals in the late 1880s, it appears that there actually was a scientific debate over the truth of this story. Naturalist C. R. Orcutt, writing in the West American Scientist in 1886, claimed to be removing all further doubt about the truth of the story by quoting an eyewitness to “the novel affair.” A writer in the bird journal Avifauna chided Spencer Fullerton Baird, secretary of the Smithsonian Institution, for perpetuating the story as fact in an article for Harper’s Magazine (Van Dyke, n.d.). I can confirm that roadrunners will attack small rattlesnakes for food and large rattlesnakes that are near a nest (see chapter 5). However, these attacks occur in reverse order from that in the thorn corral tale. Instead of being asleep at the beginning of the tale, the snake was “sleeping” in a coil after the roadrunner attack (plate 14 and the drawing at the opening of this chapter). I wonder if someone who came upon a roadrunner standing beside such a coiled snake would think that the snake was sleeping and the roadrunner was going to build a corral. Several of the eyewitness accounts retold by Dobie describe the snake being coiled as it slept.

Names for the Roadrunner The Spanish explorers who entered Mexico and the southwestern United States were the first Euro-Americans to see roadrunners. One of their names for it was “faisán real,” or “royal pheasant.” This and other Spanish names for the roadrunner are commonly used in the southwestern United States and Mexico today. Probably the two most commonly used Spanish

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names in the southwestern United States are “corre camino” (“it runs the road”) and “paisano” (“fellow countryman”). In the late 1880s and early 1900s the name “chaparral cock” appeared in most natural history writings about the roadrunner in English. This name persists today in some southwestern areas. Table 5 lists some of the colloquial names for the roadrunner.

The Roadrunner in Modern Culture Southwestern Icon The roadrunner continues to be a favorite bird and icon of the Southwest. It is the state bird of New Mexico and the emblem of the Texas Folklore Society. The roadrunner appeared often in popular articles by explorers, naturalists, and pioneers long before the animated version came along. This 1915 depiction of the roadrunner by naturalist D. I. Shepardson is a good example: “No picture of the desert is complete without him. Given a giant cactus, a rattlesnake, a lone cowpuncher and a Road-runner: and your short-story novelist has ample material for an absorbing romance of the plains” (1915, 158). Early descriptions of the roadrunner often included a humorous story or two about the bird, even coming from serious ornithologists, such as naturalist J. L. Sloanaker. In 1913 he wrote in the Wilson Bulletin, a bird journal: “Of all the birds on our list the Roadrunner is doubtless the most unique; indeed, he is queer, and would certainly take first prize in the freak class at the Arizona state fair” (Sloanaker 1913, 191). In 1940 ornithologist George Miksch Sutton wrote: “So odd, so even funny a creature is the roadrunner that it is natural to caricature him a bit in describing him” (1940, 37).

Animation of the Roadrunner Since its appearance in the 1950s, the Warner Brothers’ roadrunner and coyote cartoon characters undoubtedly have added greatly to the bird’s popularity and use as a logo in modern culture. Fondness for the animated character sparks a genuine curiosity about the real bird, which I have used to good advantage to help introduce the real bird to U.S. audiences over the last thirty years. Most people, unfamiliar with the real bird, want a comparison with the animation: Does it go “Beep! Beep!”? Does it really run only on the road? Do coyotes really chase roadrunners? How fast can they really run? I think the roadrunner’s comic reputation comes from its ungainly movements and the situations where its curiosity and boldness lead it. Indeed, there are even a few similarities between the real bird and the animation. The late great character animation film director Chuck Jones captured some of the habits of the real bird and its quirky, gutsy personality very well, and he also put a little of the roadrunner personality into the coyote character. Curious about the origin of the animated version, I interviewed Chuck Jones under the sponsorship of the National Geographic Society for an article about the roadrunner for

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Table 5 Common names for the roadrunner Type of common name

Common name

English names in the Southwest and Mexico

chaparral cock, chaparral, cock-of-the-desert, groundcuckoo, running cuckoo, racer, lizard bird, snake bird, snake-killer, war-bird, medicine bird, large roadrunner

Spanish names in the Southwest and Mexico

el correo del camino—the runner of the road or path; correr del paisano—messenger of his countryman; el paisano—compatriot or fellow countryman or peasant; la churrea or churea or churca—an imitation of its call; corre camino—it runs the road; tacó—bite, snack, or heel of a shoe (from taconar—to click one’s heels, to walk hard on one’s heels); arriero—(from arriesgar—to risk or dare); faisán real—royal pheasant

Guanajuato, Mexico

faisán—pheasant

Michoacán, Mexico

faisán mexicano—Mexican pheasant

American Indian Cochiti Pueblo (N.Mex.) Zuni Pueblo (N.Mex.) Sia Pueblo (N.Mex.) Santa Ana, Cochiti Laguna, Santa Domingo, and San Felipe Pueblos (N. Mex.) Pima (southern Ariz.) Paiute (southern Nev.) Cahuilla (southeastern Calif.) San Carlos Apache Shoshone Luiseño

sha-shua poyyi djáck´ cá ck´

d´adai soonung´wuvee toowuv puuis góshdiyé pooepooe

Sources: Simmons 1925; Dobie 1939; Lange 1959; Ladd 1963; Birkenstein and Tomlinson 1981; White 1986; Cornett 2001; Machula, n.d.

National Geographic magazine (Whitson 1983). I was pleased to learn that he based his caricature on his memories of roadrunners from when he was a lad in Southern California. He expressed a real affection for the bird, and we had quite an enjoyable time answering each other’s questions about our versions of the bird. Details of the creation and characteristics of the animated roadrunner appear in his delightful book Chuck Amuck (Jones 1999). The cartoon establishes the invincible nature of the roadrunner against the coyote, more by the coyote’s own ineptitude than by anything the roadrunner does. Ironically, this theme appears in a common folk story about the roadrunner—the corral of thorns, described earlier in this chapter. In this tale the roadrunner taunts the rattlesnake to kill itself through its own foolishness.

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Symbol of Speed The roadrunner is often used as a logo when a business wants to convey speed. Throughout the Southwest there are gas stations, truck stops, truck lines, bus companies, courier services, motels, restaurants, gift shops, and myriad other travel-related businesses that use the roadrunner as their symbol. Probably the most famous use as a speed symbol was by the auto manufacturer Plymouth, when it produced the Plymouth Roadrunner, a 1960s sports car. While maintaining captive roadrunners in Norman, Oklahoma, for my behavioral studies in the 1960s, I was approached by the local Plymouth dealer, who thought I should be thrilled to let him use my captive roadrunners in a television commercial promoting the car. Naturally, I was not interested in exposing my research animals to the trauma of a television studio. However, Dr. George Miksch Sutton graciously allowed the dealer to use one of the stuffed mounts from the Bird Range at the University of Oklahoma. As an icon for speed, the roadrunner has also been adopted as a logo by computer services, recording companies, running clubs, motorcycle clubs, hockey teams, and public schools’ sports teams, to name a few. During the Vietnam War, it was reported to be one of the favorite insignias of the U.S. soldier.

Popularity With People Were there no folklore, healing uses, cartoons, or other cultural forces at work to call attention to the roadrunner, it would still be popular with people because of its curious nature and its ready tameness. Wild roadrunners living near homes, ranches, and campgrounds are often approachable, taking food and water put out for them. There are many popular accounts of roadrunners regularly coming by a house for a handout and nesting or roosting on a window ledge or in a garage. All of this tolerance of human activity has probably helped the range expansion of the species, along with the habitat changes that we humans make in its favor (see chapter 8). The roadrunner’s tameness and curiosity provide people with a chance to get to know and love this bird. And roadrunners have provided many a humorous moment for us, such as when they fight their own reflection in a glass door or a hubcap; tease a pet dog or cat; or run-glide in front of our car, as if to taunt us. Even scientists studying the roadrunner can lose their objectivity and become quite fond of this bird. One of the first to study wild roadrunners was noted ornithologist and bird artist George Miksch Sutton. He began studying the species when only a boy of fifteen and kept several of the birds as pets (Sutton 1922). While still a teenager, he published several articles in bird journals on roadrunner behavior, as well as a drawing of one of his pets (Sutton 1913, 1915). He passed on his lifelong enthusiasm and fondness for this bird to me when I studied ornithology with him at the University of Oklahoma. In the wake of an article I wrote on the roadrunner for National Geographic magazine, I received many letters from readers who wrote about their “pet” roadrunner that was roosting,

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nesting, or feeding in their yard. With what affection they spoke about the bird! Most gave a name to their wild “pet” and put out food for it. Many reported that after the bird had been around for a while, it would take food from their hands. I don’t doubt this, as I have had the pleasure of feeding a wild roadrunner by hand too. Truly, the roadrunner is a bird of the people.

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Index

Activity patterns of roadrunner, 29–30, 87 Age determination of roadrunners, 70, 71–73, 72 (table 3), 73 (table 4) Aggressive behavior of roadrunner, 41, 46–47, 50, 56 American redstart, Setophaga ruticilla, 22 Animal trails: in roadrunner folklore 99–100; roadrunner’s use of, 21 Animation of the roadrunner, 102–103 Anis. See Groove–billed ani and Smoothbilled ani Apache Indians, 96, 103 (table 5) Aplomado falcon, Falco femoralis, 34 Arizona, 38, 39, 49, 51, 53, 60, 78, 82, 92, 96, 102 Arizona Game and Fish Department, 47, 50, 53 Arkansas, 13, 25, 39, 79, 81, 83, 84, 85, 88 Arkansas River, 84, 85 Armadillo. See Nine-banded armadillo Army ants (genus Eciton), 27 Arnold, Keith A., 11, 38

Bailey, Florence Merriam, 22, 92 Baird, Spencer Fullerton, 4, 82, 101 Banded ground-cuckoo, Neomorphus radiolosus, 8 (table 1), 14 Bandelier, Adolph, 94 Banding data for roadrunner, 14, 40 Barrel cactus (genus Ferocactus), 34 Beal, Kathleen G., 24, 33 Berger, Andrew J., 9, 10 Big Bend area of Texas, 97 Black-billed cuckoo, Coccyzus erythropthalmus: erect, posture of juvenile, 63; as rain crow, 93 Black-chinned hummingbird, Archilochis alexandri, 23 Bobcat, Lynx rufus, 34 Body fat of roadrunner: of incubating male, 55; in winter, 33 Body temperature of roadrunner: of adult, 30, 31, 32; of incubating male, 55; of nestling, 59 Breeding densities of roadrunner, 78

118

Breeding season of roadrunner: length of, 37–38; peak times of, 38; relation to food supply of, 38, 39; stages of, 38 (table 2); time of first breeding, 64. See also Renesting and Chronology Breeding success: factors in, 38; measured by number that fledge, 55 Breeding territory of roadrunner, 41, 47, 78–79 Brood parasitism: by cuckoos, 7; by roadrunner, 53 Brood patch of roadrunner, 53 Brooding by roadrunner, 59–60. See also Nestlings Bryant, Harold C., 18, 21, 77 Bull snake, Pituophis melanoleucus, 55 Cahuilla Indians, 103 (table 5) Calder, William A, 29, 30, 31, 53, 61 California, 4, 12, 39, 51, 52, 53, 56, 78, 79, 82, 83, 87, 88, 99, 103 California quail, Callipepla californica, 23 Canada, 85 Cannabalism in roadrunner, 60 Cardinal. See Northern cardinal Carotid network of roadrunner, 30 Carrion, 20, 21, 34 Cartoon roadrunner. See Animation Cascade Mountains, 87 Cat (domestic), 34 Caterpillars, 18, 22, 39, 62 Chihuahuan Desert, 51, 77, 78 Chihuahuan spotted whiptail lizard (Aspidosscelis exsanguis), Plate 9 Cholla cactus (genus Opuntia), 34, 51 Chronology of roadrunner breeding, 38, 52, 53, 62 Climate and roadrunner, 85–87 Clinal variation of roadrunner, 9 Clutch size. See Eggs

Index

Coccyzus americanus. See Yellow-billed cuckoo Coccyzus erythropthalmus. See Black-billed cuckoo Collared lizard, Crotaphytus collaris, 84 Colorado, 4, 13, 82, 83, 84, 87 Common poorwill. See Poorwill Common raven. See Raven Communal nesting by roadrunner, 53, 64 Conkling’s roadrunner, Geococcyx californianus conklingi, 7, 80 Cooper’s hawk, Accipiter cooperii, 34 Cornell Laboratory of Ornithology. See North American Nest Record Card Program Cornett, James W., 18, 32, 39, 81, 103 Coues, Elliott, 4, 82 Courtship calls of roadrunner: bark, 42, 46, 47, 50, 52, 53; clack (rattle), 42, 44, 46, 53; coo, 41, 47, 49, 50, 52, 53, 62, 65; growl, 46, 50, 52, 63, 65; whine, 46, 47, 50, 52, 62; whirr, 43 Courtship displays of roadrunner: chase, 50; prance, 42, 43, 47, 50; stick offer, 46, 47, 50, 62; tail and head flick, 45, 46; tail wag, 43, 44, 47, 50, 52, 53, 65; vertical flick, 44, 47 Courtship feeding by roadrunner: during care of fledglings, 65; during care of nestlings, 62; food exchange, 11, 45; during egg-laying, 53; during nest building, 52; during tail-wagging, 43, 45; relation to food supply, 40, 65, 80; response to coo tape, 47 Cowboy bird, 27 Coyote, Canis latrans, 34, 102, 103 Crayfish, 26 Crest and color patch: of adult roadrunner, 11–12; of nestling roadrunner, 68, 69; of lesser roadrunner, 12; in Pima Indian folklore, 97

Index

Cuckoo family (Cuculidae), 4–7; brood parasitism, 7; courtship feeding, 40 Cuculiformes order of birds, 5 Curanderos (Mexican folk-healers), 96–99, 100 Cursoricoccyx geraldinae, 7 Dark-eyed junco, Junco hyemalis, 19 Death Valley, California, 13 Desert Tortoise, Gopherus agassizii, 18, 80 Diet of roadrunner: adult, 18–26; nestling, 20–21, 25; quantities in, 21; as water source, 31; winter, 33 Directive markings of nestling, 69 Diurnal activity. See Activity pattern of roadrunner Dobie, J. Frank, 17, 92, 93, 99, 100, 103 Dormancy. See Hypothermia Dresser, H. E., 99 Drinking by roadrunner, 31 Dromococcyx pavoninus. See Pavonine cuckoo Dromococcyx phasianellus. See Pheasant cuckoo Egg tooth of roadrunner, 68 Eggs of roadrunner: clutch size, 53, 55; description of, 52; energy for laying, 55; hatching of, 53, 67–68; incubation of, 53–54; laying dates, 39; in common raven nest, 53; predation on, 35, 55; recognition of by parent, 62; staggered hatching of, 53, 59; time to lay, 52 Eltaher, Hassan, 53, 61 Endangered status of roadrunner, 14, 77 Energy conservation by roadrunner, 31, 32 European cuckoo, Cuculus canorus, 5 Eye ring of roadrunner, 70, 73 (table 4) Feathers of roadrunner: description, 9; in Pueblo Indian ceremonies, 92–95; in Anasazi art, 95

119

Fecal sacs of nestling roadrunners: 61–62, 72 (table 3); urine concentration of, 61; water source for adults, 31, 61 Feeding methods of roadrunner: beating prey, 24; handling tarantulas, 26; killing prey, 24; food pellets, 26; plucking bird feathers, 24; rolling cactus, 26; swallowing prey, 25. See also Foraging Fewkes, Jesse Walter, 93 Fledging of nestling roadrunner: by age, 62; early, 62; role of growl in, 63 Fledglings of roadrunner: courtship displays by, 65; description of, 70, 71, 73 (table 4); first breeding by, 64; foraging with parent, 63, 64; helping at the nest, 64; movement of, 63; parental attacks on, 64; feeding of, 62, 63, 64; play by, 65; response to danger, 63; sounds made by, 62–65, 73; wing-flashing by, 64 Flying behavior of roadrunner, 11 Folklore about roadrunner, 17, 91; corral of thorns, 101; Mexican, 97–101; other American Indian, 96–97; Pueblo Indian, 92–96. See also Curanderos, Crest, Feathers, Running, Superstitions, and Tracks Folse, Leon J., Jr., 11, 38, 39, 51, 52, 71, 78 Food pellets, 26, 60, 73 (table 4) Food shortages: survival of roadrunner during, 32; effect on growth of young 71 Foraging by roadrunner: at bird feeders, 23; by following deer, 23; by following humans, 23–24; in snow, 86; in tall grass, 22; at traps and mist nets, 22–23; in winter, 33; wing-flashing used in, 22. See also Feeding methods Fossils: of Conkling’s roadrunner, 7, 80; of ground-cuckoos 7; of roadrunner, 7, 9, 79–80

120

Index

Geluso, Kenneth N., 33 Geococcyx, 4, 7 Geococcyx californianus. See Greater roadrunner Geococcyx californianus californianus, 7 Geococcyx californianus conklingi. See Conkling’s roadrunner Geococcyx velox. See Lesser roadrunner Geographic distribution of roadrunner, 12–13, 83, 84, 85. See also Range expansion Gorsuch, D. M., 18, 19 Great Basin Desert, 87 Great Plains, 81 Greater roadrunner, Geococcyx californianus. See specific subject entries about roadrunner Grinnell, Joseph, 52 Groove-billed ani, Crotophaga sulcirostris juveniles assisting at nest, 64; night incubation by male 54 Ground-cuckoo/roadrunner subfamily, Neomorphinae, 5, 7, 8 (table 1); food habits, 26, 27 Ground nesting by roadrunner, 51 Gular flutter of roadrunner: adult, 30; nestling, 73, 74 Habitat of roadrunner: characteristics of, 13–14, 81; effects on distribution, 87–88; elevations of, 13, 87; fire suppression in, 88; in urban areas, 13, 79, 88; prehistoric, 79–80 Hawks, 34. See also Aplomodo falcon, Cooper’s hawk, Northern harrier and Red-tailed hawk Honey mesquite, Prosopis glandulosa, 55, Hospoa (roadrunner) kachina, 93 House finch, Carpodacus mexicanus, 23 House sparrow, Passer domesticus, 20, 23, 25

Hughes, Janice M., 9, 19, 51, 52, 70, 71, 88 Humans: affection for roadrunner by, 104–105; eating roadrunners, 93, 99; feeding roadrunners by, 20, 23, 24; 33; near roadrunner nests, 52; near roadrunner roosts, 35; persecution of roadrunners by, 14, 34, 77; providing nesting material, 52; role in roadrunner expansion, 80–85, 88. See also Folklore and Range Expansion Hummingbirds: aerial wing-popping, 42; in roadrunner diet 23. See also Blackchinned hummingbird Hypothermia: of roadrunner, 31, 32, 33, 35, 54, 55, 86; of other birds, 33 Inca dove courtship display, 44 Incubation. See Eggs Iowa, 85 Jaeger, Edmund C., 21, 33 Jones, Chuck, 102–103 Juvenile roadrunners. See Fledglings Kansas, 77, 79, 81–84, 88 Kartchner Caverns State Park, Arizona, 9 Kidney of roadrunner, 31 Kit fox, Vulper macrotis, 34 Klamath Mountains, 87 Ladd, Edmund J, 92, 93, 103 Leopold, Aldo, 17 Lesser ground-cuckoo, Morococcyx erythropygus, 8 (table 1); following army ant swarms, 27 Lesser roadrunner, Geococcyx velox, 4, 7, 8 (table 1); appearance, 12; diet, 26; geographic range, 13 Lesson, René Primèvere, 4

Index

Lizards, 18, 20, 21, 24, 25, 26, 40, 45, 47, 60, 62, 63, 84, 97, 101. See also Collared lizard and Chihuahuan spotted whiptail Longevity of roadrunner, 14 Louisiana, 13, 79, 81, 84, 85, 88 Madrid, Enrique, Jr., 97 Marshall, Joe T., Jr., 88 Massasauga rattlesnake, Sistrurus catenatus, 20 Mate-selection by roadrunner. See Courtship and Pair-bonding Meinzer, Wyman, 14, 19, 20, 22, 25, 26, 34, 40 Mexico, 12, 13, 79, 80, 82, 87, 88, 93, 96, 97–100, 103 Migration of roadrunner, 12, 13 Mississippi, 85 Mississippi River, 79, 84, 85, 88 Missouri, 13, 14, 79, 84, 85, 88 Missouri River, 84, 85 Mojave Desert, 78 Molt of roadrunner: adult, 71; first, 70 Monkeys (tamarins, genus Saguinas), 27 Morococcyx erythropygus. See Lesser ground-cuckoo Mountain lion, 95–96 Movement of roadrunner: during lifetime, 40; on overcast days, 87. See also Banding data and Range expansion Muller, Kerry A., 71, 72, 73 Names for the roadrunner, 101–102; 103 (table 5) Nasal salt gland: of adult roadrunner, 30, 31, 32; of nestling roadrunner, 31, 32, 72 (table 3), 74; of other birds, 31, 74 National Zoological Park, Washington, D. C., 72

121

Neomophinae. See Ground-cuckoo/roadrunner subfamily Neomorphus, 7 Neomorphus geoffroyi. See Rufous-vented ground-cuckoo Neomorphus pucheranii. See Red-bellied ground-cuckoo Neomorphus radiolosus. See Banded ground-cuckoo Neomorphus rufipennis. See Rufouswinged ground-cuckoo Neomorphus squamiger. See Scaled groundcuckoo Nesting success. See Breeding success Nestlings of roadrunner: begging by, 60, 68–69; development of, 68–69, 71, 72 (table 3); diet of, 20–21, 25, 60; food shortages, 60–61; movement by, 69–70; parental care of, 59–62; predation on, 35, 55–56; sounds of, 42, 60, 68, 69, 70; temperature control in, 59, 71, 73–74; voiding waste by, 61. See also Nasal salt gland Nests of roadrunner: building 50, 62; description, 52; heights, 51; open areas around, 51; sun orientation of, 51; protection of, 55–56; removing waste from, 61–62; selection of sites for, 50–52; types of supports for, 51–52. See also Renesting Nevada, 12, 82, 87 New Mexico, 39, 49, 51, 77, 78, 79, 83, 87, 92–96, 102 Newberry, John, S., 83, 99 Nice, Margaret Morse, 83 Nine-banded armadillo, Dasypus novemcinctus, 88–89 North American Breeding Bird Survey, 14 North American Nest Record Card Program, 49

122

Index

Northern bobwhite quail, Colinus virginianus, 19 Northern cardinal, Cardinalis cardinalis, 19 Northern harrier (hawk), Circus cyaneus, 34 Northern mockingbird, Mimus polyglottos: wing-flashing by, 22, 24; roadrunner egg in nest of, 53 Nuptial feeding. See Courtship feeding Oberholser, Harry C., 9, 39, 53, 100 Ohmart, Robert D., 20, 31, 32, 38, 51, 60, 71–74 Oklahoma, 31, 32, 33, 49, 67, 70, 74, 77, 79, 81, 83, 84, 88 Orchard oriole, Icterus spurius, 19 Oregon, 85, 87 Oregon junco. See Dark-eyed junco Pair-bonding of roadrunner, 39–40, 49–50 Paiute Indians, 103 (table 5) Parental behavior of roadrunner. See Brooding, Eggs, Nestlings, Fledging, and Fledglings Parsons, Elsie Clews, 92, 94 Patuxent Wildlife Research Center: birdbanding data, 14, 40; stomach content data, 18, 19, 21 Pavonine cuckoo, Dromococcyz pavoninus, 8 (table 1) Peccary bird, 27 Pellets. See Food pellets Persecution of roadrunner. See Humans Pet roadrunners, 99, 104–105 Pheasant cuckoo, Dromococcyx phasianellus, 8, 22; diet of, 26, 27; wingflashing by, 22, 27 Physical features of adult roadrunner, 9–12. See also Crest and color patch Physical features of young roadrunner: fledglings, 70–71, 73 (table 4); nestlings, 68–69, 72 (table 3)

Pike, Zebulon Montgomery, 4 Pima Indians, 97, 103 (table 5) Poorwill (Common), Phalaenoptilus nuttallii, 33 Population of roadrunner, 14, 77–79; frequencies, 77–78; densities, 78 Pottery Mound, New Mexico, 95 Predation: defending nest from, 55–56; erect posture of juvenile roadrunner to, 63; protection from, 34–35, 51, 53, 55, 61; on roadrunners, 34, 55, 77; roadrunner vulnerable to, 25, 31–32 Prickly-pear cactus (Opuntia species), 19, 23, 51, 62 Protected status of roadrunner, 14, 77 Pueblo Indians, 92–95, 103 (table 5) Quail: egg in roadrunner nest, 53; killing of by roadrunner, 17, 18, 19. See also California quail and Northern bobwhite quail Rain crow, 93, 100 Range expansion of roadrunner, 79–85; barriers to, 85–89; factors influencing, 80–81; humans and, 80–85, 88; history of, 81–85 Raven (common), Corvus corax, 53 Red-billed ground-cuckoo, Neomorphus pucheranii, 8 (table 1); diet of, 27; following army ant swarms, 27; following monkey troops, 27 Red-tailed hawk, Buteo jamaicensis, attacking roadrunner, 34; roadrunner in stomach of, 34 Renesting of roadrunner, 37, 38, 62, 95 Research needs: courtship food, 21, 40; diet in northern and eastern range, 21; digestion of vertebrates by nestlings, 26; ecology in northern and eastern range, 85; food pellets, 26; life-time pair bonding,

Index

39; nestling salt gland, 74; sexual dimorphism, 11; winter survival, 33 Rio Grande, 88, 92, 99 Riverbanks Zoological Park, Colombia, South Carolina, 39 Roadrunner as symbol: for Southwest, 91, 102; for speed, 104 Rocky Mountains, 13 Roosting of roadrunner, 34, 35, 50 Root, Terry, L., 85–87 Round-tailed horned lizard, Phrynosoma modestum, 25 Rufous-vented ground-cuckoo, Neomorphus geoffroyi, 6, 7, 8 (table 1); diet of, 26, 27; following army ant swarms, 27; following monkey troops, 27; nest height of, 51 Rufous-winged ground-cuckoo, Neomorphus rufipennis, 8 (table 1); following peccaries, 27 Running ability of roadrunner, 10; running speeds, 10–11; in folklore, 93, 96–97 Salt gland. See Nasal salt gland Sand bathing of roadrunner, 32, 70 Saurothera, 4 Scaled ground-cuckoo, Neomorphus squamiger, 8, 14 Schaafsma, Polly, 93, 95–96 Scorpions, 17, 18, 21, 26, 29, 98 Sexual dimorphism of roadrunner, 11 Shading behavior of roadrunner: in heat, 30; of nestlings, 59 Shepardson, D. I., 102 Sherbrooke, Wade C., 25, 56 Shoshone Luiseño Indians, 103 (table 5) Shufeldt, R. W., 10 Sierra Nevada, 87 Skutch, Alexander F., 26, 27, 53 Sloanaker, J. L., 102 Smithers, W. D., 98

123

Smooth-billed ani, Crotophaga ani: communal nesting, 53; night incubation by male, 54 Snails, 18, 26 Snake: as egg predator, 49, 55; attack of by roadrunner, 19, 24, 25, 34, 55, 56; in roadrunner diet, 18–19, in folklore, 92, 98, 101 Snakebird, 18 Sonoran Desert, 38, 51, 78 Sounds of roadrunner. See Courtship calls, Nestlings, and Fledglings Spofford, Sally Hoyt, 23 Spruce grouse, Falcipennis canadensis, wing-snapping display, 42 Stevenson, M. C., 92 Stomach content studies, 18–19, 26; winter, 33 Striped cuckoo, Tapera naevia, 8 (table 1); alula-flashing, 27; tail-wagging display, 44 Subspecies of roadrunner, 9 Sumac (genus Rhus), 19 Sunning behavior: of adult roadrunner, 32, 55, 70; of nestling roadrunner, 70; of other birds, 32 Superstitions about roadrunners, 99–100 Survival mechanisms: in cold weather, 31–32; in dry conditions, 31; in food shortages, 32–34; in hot weather, 29–30 Sutton, George Miksch, 17, 32 , 34, 52, 53, 63, 87, 102, 104 Sweat glands, 30 Tapera naevia. See Striped cuckoo Tarahumara Indians, 93 Tarantulas, 17, 26, 98 Tasajillo cactus, Opuntia leptocaulis, 26 Temperature control of roadrunners, 30, 31, 32-33; in cold climates, 87; during incubation, 55; of nestlings, 59, 71, 73–74

124

Index

Territoriality of roadrunner, 39, 41, 47, 49, 50, 56; See also Aggressive behavior and Breeding territory Texas, 3, 25, 38, 49, 51-54, 60, 62, 74, 77, 78, 81, 83, 84, 88, 95, 97, 102 Texas horned lizard, Phrynosoma cornutum, 20, 25 Threatened status of roadrunner, 14, 77 Torpor: of poorwill, 33; of roadrunner, 3233, 86; of white-throated swift, 33 Tracks of roadrunner, 94, 100; in petroglyphs 96; use by Pueblo Indians, 94; use by Anasazi, 95–96 Tyler, Hamilton A., 92, 95 Urban roadrunners, 13, 20, 34, 79, 81, 88 Urine concentration: of adult roadrunners, 31; of mammals, 31; of nestling roadrunners, 61, 67, 74 Utah, 12, 83, 84, 87 Vehrencamp, Sandra L., 54, 55 Water balance: adult roadrunners, 30, 31, 61; nestling roadrunners, 74 Welder Wildlife Refuge, Texas, 78 Western coral snake, Micruroides euryxanthus, 19

Western diamondback rattlesnake, Crotalus atrox, 19, 56, White-tailed deer, Odocoileus virginianus, 23 White-throated swift, Aeronautes saxatalis, 33 Wing-flashing: by adult roadrunner, 22, 23, 24, 56; by ground-cuckoos, 22, 27; by juvenile roadrunner, 22, 64; by other birds 22 Winter survival of roadrunner: and body fat, 33; 32–34; and effect on distribution, 85–87; and food, 18; and foraging, 33; and shelter, 33 Wright, Robert, E., 26 Yellow mud turtle, Kinosternon flavescens, 18 Yellow-billed cuckoo, Coccyzus americanus, 5; breeding season of, 39; coo call of, 41; female tail-up posture of, 44; longevity of, 14; as rain crow, 93, 100; stick offering by, 46 Yucca (genus Yucca), 24, 55, 94 Zimmerman, Dale A., 23 Zygodactyl foot, 6, 9, 72 (table 3)