Redacted for privacy Abstract Approved:

AN ABSTRACT OF THE THESIS OF JAMES DALE STEPHENSON for the MASTER OF SCIENCE (Name) (Degree) in WILDLIFE SCIENCE presented on August 27, 1970 (Major) (Date) Title: PLUMAGE DEVELOPMENT AND GROWTH OF YOUNG WHOOPING CRANES Redacted for privacy Abstract Approved: Holiard M. Wight Plumage development and growth were studied in young whooping cranes (Grus americana) from hatching until 450 days of age. The downy stage of the young cranes lasted about six weeks; the development of the juvenal plumage took until 100 days of age, and the molt to the adult or all white plumage was complete at 450 days. At one day the cranes stood 19.0 to 20.5 centimeters in height and weighed 100 to 150 grams. By 40 days they stood 65.0 to 75.0 centimeters and weighed 1,750 to 2,500 grams; by 100 days they stood 100 to 120 centimeters and weighed 4,300 to 4,950 grams; and by 250 days they were nearing adult size and stood 112 to 127 centimeters and weighed 5,700 to 6,700 grams. The weights for each of six cranes were graphically plotted, and a logistic growth curve was determined for each individual. A mean logistic growth curve was computed from the accumulated data. Color changes of the down, contour feathers and other body parts were noted and a key to age identification based on these observations was developed.

Plumage Development and Growth of Young Whooping Cranes by James Dale Stephenson A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science June 1971

APPROVED: Redacted for privacy Associate Professor of Wildlag/Ecology Redacted for privacy Head of Department of Fisheries and Wildlife Redacted for privacy Dean of Graduate School Date thesis is presented August 27, 1970 Typed by Carolyn Mills for James Dale Stephenson

ACKNOWLEDGMENTS I wish to thank Ray C. Erickson for suggesting the study and offering direction and encouragement. I also wish to thank other members of the staff at the Patuxent Wildlife Research Center who offered their encouragement and advice. Special appreciation is extended to Robert G. Heath for his assistance in analyzing the growth data. My wife, Mary, deserves many thanks for her help in gathering data and criticism of the manuscript. Howard M. Wight, my major professor, provided many helpful suggestions to improve the manuscript.

TABLE OF CONTENTS Page INTRODUCTION MATERIALS AND METHODS RESULTS 1 2 4 The Newly Hatched Chick 4 One Day 4 Five Days 5 Eleven Days 6 Eighteen Days 6 Twenty-eight Days 6 Forty Days 7 Forty-nine Days 8 Seventy-five Days 9 Ninety-five Days 10 One Hundred Twenty-five Days 10 One Hundred Fifty Days 11 One Hundred Seventy-five Days 11 Two Hundred Days 12 Two Hundred Fifty Days 12 Three Hundred Days 12 Three Hundred Fifty Days 13 Four Hundred Days 13 Four Hundred Fifty Days 13 Anatomical Measurements 31 Height 31 Weight 31 DISCUSSION 43 The Downy Stage 43 Development of the Juvenal Plumage 43 The Juvenal Molt 44 Height 45 Weight 45 KEY TO THE AGE OF YOUNG WHOOPING CRANES 51 LITERATURE CITED 53 APPENDIX I 54

LIST OF FIGURES Figure Page 1. One day. 14 2. Five days. 15 3. Eleven days. 16 4. Eighteen days. 17 5. Twenty-eight days. 18 6. Forty days. 19 7. Forty-nine days. 20 8. Seventy-five days. 21 9. Ninety-five days. 22 10. Left: Two days. Right: Forty days. 23 11. Left: Fifty days. Right: Seventy-five days. 24 12. Left: One hundred days. Right: One hundred fifty days. 25 13. Left: Three hundred days. Right: Adult. 26 14. Left: Seventy-five days. Right: One hundred days. 27 15. Left: Two hundred days. Right: Two hundred fifty days. 28 16. Left: Three hundred fifty days. Right: Adult. 29 17. One hundred twenty-five days. 30 18. Heights of six whooping cranes. 34 19. Logistic growth curve for crane 68-2. 36 20. Logistic growth curve for crane 68-3. 37 21. Logistic growth curve for crane 68-4. 38 22. Logistic growth curve for crane 68-6. 39

Figure Page 23. Logistic growth curve for crane 68-7. 40 24. Logistic growth curve for crane 68-8. 41 25. Mean logistic growth curve for six whooping cranes. 42

LIST OF TABLES Table I. Whooping crane chick measurements Page 33 II. Logistic growth curve values and means for weights of six whooping cranes. 35 III. Progressive changes in appearance useful in identifying age classes. 48

Plumage Development and Growth of Young Whooping Cranes INTRODUCTION The major purpose of this study was to provide detailed information on the progressive development of whooping cranes (Grus americana) of known age. literature. This information was not previously available in the These data consist of color descriptions of down, contour feathers and other body parts, replacement of plumage, weights, heights, and other physical changes. Limited observations of growth and plumage development in whooping cranes had been made by other authors. Blyth and Tegetmeier (1881) briefly describe adult and immature cranes, but not the downy young. Bent (1926) gives Bradshaw's descriptions of the downy young and describes young-of-the-year taken in October and February. Adult and juvenal plumages and natal down are also described by Ridgway and Friedmann (1941). Stevenson and Griffith (1946) noted the progressive changes in appearance of the young on the wintering grounds, and Allen (1952) devotes a small section of his monograph to molts and plumages, quoting some of the above authors.

2 MATERIALS AND METHODS This study was based on captive birds of known age. In 1967, 1968, and 1969, eggs were collected from the whooping crane's breeding grounds in northern Canada and brought to Patuxent Wildlife Research Center near Laurel, Maryland, where incubation was completed in mechanical incubators. Five birds were hatched from six eggs in 1967 and another chick was hatched at the San Antonio Zoo that year and brought to Patuxent. Ten birds hatched from ten eggs in 1968, while seven birds hatched from ten eggs in 1969. Most of the data were collected from the 1968 birds, although some data were taken from the 1967 birds between January 1968 and June 1968. Material was also gathered from the 1969 hatched birds for comparative purposes during June 1969. Much of the data collected on plumage development and color changes were recorded on duplicated line drawings of the birds. Four different drawings were made to account for the progressive changes in body proportions as the birds grew. All colors described are those taken from Ridgway's Color Standards and Color Nomenclature (1912). These are listed in the appendix beside the corresponding plate number from Ridgway. Feather tracts (pterylae) mentioned in the text are those presented by Beister and Schwarte (1965). Measurements were taken of the culmen, bill depth, tarsus, and middle toe from seven of the birds hatched in 1968 when they were one to seven days of age. The measurements were made using a small set of

3 calipers. To avoid the risk of injuring the birds, no further anatomical measurements were taken as the birds grew. Weights were taken daily through the first two to three weeks of age, weekly until the end of December, and bimonthly thereafter. The cranes were initially fed a diet of commercial poultry feed, and mealworms (Tenebrio molatar). It was necessary to hand feed the birds moistened feed for the first few days. The mealworms were added as an incentive for the birds to eat on their own, and were discontinued soon thereafter. The commercial poultry feed, a feeder and maintainer diet, was used throughout the course of the study. During the first two weeks, estimates of heights were made as the birds stood in front of a calibrated section of plywood that was part of the pen. After the birds were moved to outdoor runs, heights were estimated as they stood in a normal position near a post or plywood panel situated in the pen. A point was thus determined even with the top of the bird's head, and measurements were made from that point to ground level with a ruler or tape. During the first month after hatching 35 mm. photographs were made at intervals of three to four days, then weekly until three months of age, and thereafter at two to three week intervals. These photographs were used later in the analysis of plumage development.

4 RESULTS The Newly Hatched Chick The newly hatched chick has swollen legs and feet with the blood vessels showing beneath the skin, which gives them a reddish cast. The chick is wet and appears dark at first, but becomes lighter in appearance as the down dries. A whitish flaky material initially ensheathes the down but is gone by the second day. Usually the chick does not try to stand for the first few hours, and is seen lying down or resting on its heels (hocks). One Day (Figure 1, Figure 10) The chicks stand 19.0 to 20.5 centimeters in height and weigh 100 to 150 grams at this age. The tip of the bill is light drab followed by a narrow band of cinnamon-brown with a narrow fringe of light drab immediately behind this on the upper mandible. The proximal two-thirds of the bill are light salmon-orange. A pale pinkish cinnamon egg tooth is present on the upper mandible. A narrow strip of mummy brown down starts at the base of the bill, extends up the center of the forehead, and forms a "Y", the branches of which terminate on the sides of the head very near and in front on the ear tuft. Cinnamon-brown covers the crown and back of the head. The eye is dark olive. Clay color encircles the eye, while pale pinkish cinnamon borders on the edge of the bill, extends onto the chin and throat, encircles the ear tuft and continues down the fore neck onto the breast. Cinnamon-brown covers

5 the hind neck and scapular area, extending down the dorsal surface of the wing and onto the flanks and legs. Pale pinkish cinnamon appears on the underside and dorsal inside corner of the wing. Sepia covers the interscapular region and blends into mummy brown which extends down the remainder of the back, across the rump and onto the dorsal surface of the tail. Pale pinkish cinnamon extends between the legs from the breast and covers the abdominal region. The tibia and tarsus, as well as the dorsal surfaces of the toes and nails, are dull purplish black. The heels, and ventral surfaces and proximal joints of the toes are light purplish vinaceous. At this stage the toenails are essentially the same color as the dorsal surfaces of the toes. Few observations were made of the toenails as the birds grew, since many times they were covered with mud or were not visible in the grass. Five Days (Figure 2) The tip of the bill has become drab instead of light drab, and the narrow band of cinnamon-brown near the tip of the bill is benzo brown. Orange-pink replaces the light salmon-orange on the proximal two-thirds of the bill. The nostrils are clay color. The eye is olivaceous black with a glassy, bluish cast. The tibia and tarsus as well as the dorsal surfaces of the toes and nails are dull violet-black. The ventral surfaces and proximal joints of the toes and the heels (hocks) are light vinaceous-fawn to light vinaceous. The birds stand approximately 25.5 centimeters in height, and weigh 120 to 165 grams.

6 Eleven Days (Figure 3) The tip of the bill is drab-gray, the narrow band of benzo brown has disappeared, and the proximal portion of the bill has changed to light ochraceous-salmon. Light pinkish cinnamon has replaced pale pinkish cinnamon on the fore neck, breast and abdominal region, and also appears in the scapular area, as well as the area encircling the eye that was previously clay color. The birds stand approximately 30.5 centimeters in height and weigh 200 to 300 grams. Eighteen Days (Figure 4) A portion of the "Y" on the forehead persists, but is cinnamonbrown in color. Cinnamon-buff is present in the area encircling the eye, with cinnamon-brown in the scapular region. A "V" of pale pinkish cinnamon appears on the breast. The birds are 33.0 to 40.5 centimeters in height and weigh 350 to 625 grams. The neck and legs have begun to lengthen, giving a much gawkier appearance. Twenty-eight Days (Figure 5) The very tip of the bill is sayal brown, followed by drab gray, with the proximal portion of the bill remaining light ochraceous-salmon. The cinnamon-brown "Y" of the forehead is now only a median strip. The eye has turned from olivaceous black to olive. A downy pale pinkish cinnamon ear tuft approximately one-fourth inch long is very noticeable. Quite a bit of dull violet-black is noticeable at the base

7 of the down with the exception of the crown, which gives the cinnamonbrown of the back the effect of sepia. The down is one-third inch long in the scapular area, one-fourth inch in length on the trailing edge of the wing, (this longer down of the wing is cinnamon-brown instead of cinnamon), and one-third to one-half inches long on the tail. A much larger area of the heels (hocks), and the ventral surfaces and proximal joints of the toes are now pale salmon color which blends into the remaining dull violet-black. The birds stand 48.0 to 53.0 centimeters in height and weigh 850 to 1,050 grams. Forty Days (Figure 6, Figure 10) The very tip of the bill has changed to grayish olive, blending into light pinkish cinnamon with a basal border of seashell pink. The cinnamon-brown median strip on the forehead has disappeared and the forehead, cheek, chin and throat are light pinkish cinnamon. The eye is dark grayish olive. Cinnamon-buff covers the crown and back of the head. The ear tuft has changed to pale pinkish cinnamon. The upper back and interscapular area is chaetura drab mixed with cinnamon. Light vinaceous-cinnamon is present in the scapular area. drab appears on the distal portion of the last wing joint. Chaetura The underlining of the wing is pale pinkish cinnamon, and the outer portion of the down covering the legs is cinnamon-buff with an inner portion of pale pinkish buff. Light vinaceous-cinnamon appears on the heels (hocks), hind portions of the tibia and tarsus, and ventral surfaces and proximal joints of the toes. The fore parts of the tibia and

8 tarsus are chaetura drab, and the dorsal surfaces of the toes are chaetura black. At this age the chicks are still entirely downy except for the emergence of the black primaries in the form of pinfeathers. Throughout the period of growth, there is a progressive lengthening of the down; more so in some areas than others (see 28 days). They stand 65.0 to 75.0 centimeters in height and weigh 1,750 to 2,500 grams. Forty-nine Days (Figure 7, Figure 11) The basal portion of the bill is pale flesh color. Black hairlike feathers have appeared along the border of the upper mandible and on the chin along the lower mandible. Cinnamon-buff encompasses the forehead, cheek, chin and throat, and a narrow band of clay color separates the forehead from the crown. The neck is cinnamon with small tufts of pale pinkish buff appearing on the upper part of the hind neck. Cinnamon-brown contour feathers cover the dorsal and interscapular tracts; cinnamon-brown contour feathers with white bases are present in the scapular area, and white based cinnamon coverts cover the remainder of the humeral tract. Downy tipped cinnamon secondaries with white bases are beginning to appear and the black primaries, primary coverts, and alula are evident. Specks of chaetura drab appear on the ends of the down covering the rump. White-based cinnamon retrices with long downy tips are beginning to emerge. Light pinkish cinnamon now covers the abdominal region, legs and breast. Pale pinkish buff appears on the fore part of the tarsus. Since about the forty-second day and the

first emergence of contour feathers, there has been a very dramatic change leading to a plumage that is half contour feathers and half down. At this age the birds stand 80.0 to 98.0 centimeters and weigh 2,450 to 3,350 grams. Seventy-five Days (Figure 8, Figure 11, Figure 14) The proximal one-third of the bill is light grayish olive followed by light ochraceous-buff with a base of flesh color. The capital tract is covered with sayal brown contour feathers, with a crest of cinnamon-buff down on the crown. The eye is light olive-gray. Rather fluffy, pinkish cinnamon contour feathers appear in the dorsal cervical and ventral cervical tracts, with white bases evident on the ventral cervical tract feathers. The scapulars have lightened to cinnamon and are edged with a very light touch of black. White marginal coverts are evident when the wings are extended. The remaining coverts form an alternating pattern of white and cinnamon, the tips being cinnamon and the bases white. The legs remain downy and are nearly white with some cinnamon and a hint of black on the ends of some of the down. The pale pinkish buff on the heels deepens to pinkish buff which blends into the chaetura black on the fore part of the tarsus. The white retrices are nearly three inches in length and have a downy cinnamon tip. The decurved cinnamon tertials are elongated and extend down onto the tail. The birds stand 100 to 115 centimeters and weigh 3,950 to 4,650 grams.

10 Ninety-five Days (Figure 9, Figure 12, Figure 14) The mid portion of the bill has changed from light ochraceous-buff to pale ochraceous-buff. The eye is smoke gray and a very light scattering of black hairlike feathers is visible on the cheek beneath the eye. The feathers on the back and scapular region on one bird have lengthened to the point where they give an effect of spotting due to the visibility of the white bases of these feathers. This pattern of white and cinnamon is absent on the remaining five birds. The abdomen is almost entirely white and there is a nearly white diamond-shaped area appearing on the breast. The leg is feathered and is nearly white but still has a tinge of cinnamon and black. These feathers have a rather fluffy appearance. The tibia, tarsus and dorsal surfaces of the toes are chaetura black with pinkish buff appearing between the scales and in two small areas on each side and below the heels (hocks). There is a very light colored crane, and one that is somewhat darker than the others. The darker individual has only a very small area of the breast which is turning lighter, and has a considerable amount of cinnamon left on the legs. The lighter bird has a much larger area of white on the breast and is generally lighter overall. The birds at this age are completely feathered and have full juvenal plumage. They stand 100 to 120 centimeters in height and weigh 4,300 to 4,950 grams. One Hundred Twenty-five Days (Figure 17) The proximal two-thirds of the bill have become grayish olive with some blending into deep grayish olive on the tip. The eye is olive

11 buff. The black is more pronounced on the cheeks and extends up onto the forehead. A few of the cinnamon feathers have been replaced by white feathers on the ventral portion of the neck. replaced on the upper back (from the dorsal tract). Some have also been The plumage across the rump and the tail coverts is nearly white, except for the cinnamon tips on the tail coverts. The tip and the margins of the outer web of the last four secondaries are black and at least half of the remaining secondaries contain some black near the tips. The tibia, tarsus, and dorsal surfaces of the feet are blackish slate as in the adults. (No pinkish buff areas remain between the scales). The birds weigh 4,825 to 5,700 grams. One Hundred Fifty Days (Figure 12) The tip of the bill is benzo brown. Immediately behind the nostrils is a small area of black which is very pronounced in one individual and is also present to a lesser degree on all of the other birds except one. The black of the cheek and forehead is very noticeable. Nearly half of the feathers have been replaced on the upper back of most of the birds. There has also been an increase in the replacement of feathers on the lower half of the neck and breast. At this time the birds weigh 5,200 to 6,075 grams. One Hundred Seventy-five Days The tip of the bill is still benzo brown, but fades into dark olive-gray, which lightens to yellowish olive to light yellowish olive.

12 In one individual the entire tip of the bill is dark olive gray. Carrot red is beginning to show through on the forehead, fading out on the edges to flesh color. The eye is cream color. The replacement of feathers on the upper neck gives a salt and pepper effect. The birds weigh 5,500 to 6,400 grams. Two Hundred Days (Figure 15) The black on the face is more concentrated on the cheek and is becoming visible on the crown. Some of the head plumage is being replaced giving a white flecked appearance to the head and neck. upper back has only a scattering of cinnamon feathers remaining. The The birds weigh 5,700 to 6,700 grams and stand 112 to 127 centimeters. Two Hundred Fifty Days (Figure 15) A large whitish patch is evident behind and below the eye bordering on the upper portion of the black cheek patch. Two of the birds have molted three or four middle wing coverts on the right wing. None of the other birds has started to molt these coverts. Three Hundred Days (Figure 13) A small area on the very tip of the upper mandible is white. Some scattered glossy black feathers (not the hairlike feathers) are present on the rear portion of the crown and back of the head. The eye is massicot yellow. the upper neck. More than half of the feathers have been replaced on The back and scapulars are completely white, and

13 three-fourths of the middle coverts have been replaced on two of the birds, one-half on another bird, one-quarter on two other birds, and none on the remaining bird. Three Hundred Fifty Days (Figure 16) The tip of the bill is light cinnamon-drab, immediately followed by light grayish olive, then grayish olive with deep grayish olive in the region of the nostrils. The nostrils are pale flesh color which extends toward the base where it blends into flesh color. Carnelian red covers the forehead and crown, and flesh color appears on the cheek. The black hairlike feathers are distributed over all of the red areas. A scattering of cinnamon persists on the upper neck and head. The majority of wing coverts have been replaced and the tertials are still cinnamon tipped. Four Hundred Days The head and neck are completely white but a few cinnamon wing coverts remain. Four Hundred Fifty Days (Figure 13, Figure 16) Full adult plumage.

14 light salmon-orange mummy brown dark olive light drab pale pinkish cinnamon. cinnamonbrown clay color 1 sepia pale pinkish cinnamon mummy brown 1 4 fr le cinnamon-/ pale pinkish cinnamon 1 \/r/f I 7-- dull purplish black. light purplish vinaceous Figure 1. One Day

15 mummy brown clay color clay color benzo brown drab e / I I olivaceous black (with a bluish cast) / orange- pink e i cinnamon-brown 1 - pale pinkish cinnamon mummy brown / / if re. /04.----pale pinkish cinnamon light grayish vinaceous dull violet black Figure 2. Five Days

16 olivaceous black mummy brown drab-gray light pinkish (,- A\ I cinnamon cinnamon light pinkish cinnamon light pinkish cinnamon pale pinkish cinnamon light ochraceoussalmon cinnamon I.,' cinnamon- / brown light pinkish cinnamon dull violet black j light grayish vinaceous Figure 3. Eleven Days

17 cinnamon-brown olivaceous black drab gray clay color cinnamon-buff light ochraceoussalmon light pinkish cinnamon N,"4--'7.- cinnamon-brown pale pinkish cinnamon cinnamon \ k \ k \ 4%, light pinkish cinnamon dull violet-black light grayish vinaceous Figure 4. Eighteen Days

18 cinnamon-brown cinnamon-buff sayal brown clay color olive drab-gray pale pinkish cinnamon (ear tuft) light pinkish cinnamon N 1 k cinnamon-brown with \ k \. dull violet-black.. beneath, giving the \ -... "%... effect of sepia \ % %-_,) \ 1 1\ \ \ I \ 1 \\\ 1.-- i... ;:...--!... i 1 % ) \ S light ochraceoussalmon cinnamonbrown down 1 '! i cinnamon \\ light pinkish cinnamon dull violet-black pale salmon color Figure 5. Twenty-eight Days

19 seashell pink cinnamon-buff dark grayish olive grayish olive light pinkish cinnamon light pinkish cinnamon chaetura drab i chaetura drab mixed with light pinkish cinnamon \ \ \ \ \ cinnamon 1 t,...-' chaetura drab mixed cinnamon \N with \ cinnamon --- light vinaceouscinnamon ''N.2...!..4...NZ.- om`l chaetura drab. ---... cinnamon -.......-....,,,,,---....- ") (------. light pinkish i cinnamon cinnamon-buff light vinaceouscinnamon chaetura drab Figure 6. Forty Days

20 clay color black, "whiskers" r grayish olive pale flesh color cinnamon-buff deep grayish olive pale pinkish buff light pinkish cinnamon small tufts of pale pinkish buff cinnamon cinnamon-brown white N cinnamonbuff cinnamonbrown white cinnamon light pinkish cinnamon light vinaceouscinnamon cinnamon Ispecks of chaetura drab (under wing on back and center lower back) on the end of the down chaetura black pale pinkish buff Figure 7. Forty-nine Days

21 flesh color cinnamon-buff (down) light \ sayal brown light olive-gray grayish 1 olive light ochraceous- / buff (/)11 I pinkish cinnamon cinnamon pinkish cinnamon with white showing at the base of the feathers \ white alternating with cinnamon black edited feathers V mostly white tinged with cinnamon and some black 1 white pale pinkish buff cinnamon (down) pinkish buff chaetura black Figure 8. Seventy-five Days

22 light grayish olive flesh color light ochraceousbuff sayal brown smoke gray black hairlike feathers (barely visible) pinkish cinnamon pinkish cinnamon with white showing at the base of the feathers cinnamon and white pattern nearly white with barely a bit of cinnamon showing white tinged with cinnamon and some black white chaetura black (pinkish buff between the scales) cinnamon (down) pinkish buff area on each side of the legs Figure 9. Ninety-five Days

Figure 10. Left: Two days. Right: Forty days.

Figure 11. Left: Fifty days. Right: Seventy-five days.

Figure 12. Left: One hundred days. Right: One hundred fifty days.

Figure 13. Left: Three hundred days. Right: Adult.

Figure 14. Left: Seventy-five days. Right: One hundred days.

Figure 15. Left: Two hundred days. Right: Two hundred fifty days.

Figure 16. Left: Three hundred fifty days. Right: Adult.

Figure 17. One hundred twenty-five days.

31 Anatomical Measurements Measurements taken during the first week of age of the culmen, bill depth, tarsus, and middle toe of seven of the birds hatched in 1968 are given in Table I. Height Height increments for the period of growth of six cranes hatched in 1968 are illustrated in figure 18. (Three of the ten birds hatched in 1968 died and one developed wing abnormalities. These birds were excluded in the height and weight data.) Weight The weights for each of six cranes hatched in 1968 were plotted, and a logistic growth curve derived from the differential equation dw = dt by (k-y) was determined for each individual. The formula for this curve [see Kendall and Buckland (1966) pages 123 and 124] is written: Y K 1 + A exp - bt where Y = wt. at time t K = final wt. asymptote A = YK-Yo o where Yo = initial wt. b = constant related to the rate of growth tj = Di-3, 0 < j < 15 Di = day of age, 3 < i < 228

32 The six cranes were identified as 68-2, 68-3, 68-4, 68-6, 68-7, and 68-8 and the data for each crane are plotted on figures 19 through 24 respectively. A mean logistic growth curve for the six birds is shown in figure 25. Table II shows the determined values for each bird and the mean of each value.

33 TABLE I. WHOOPING CRANE CHICK MEASUREMENTS Bird No. Abe Culmen Bill Depth Tarsus Middle Toe 68-1 1 day 24.1 mm 12.6 mm 45.8 mm 32.9 mm 68-2 2 days 24.6 12.3 51.2 37.1 68-3 7 days 29.0 14.0 57.0 39.9 68-4 3 days 24.5 12.3 49.5 35.5 68-8 3 days 25.8 13.4 52.4 35.0 68-9 7 days 28.5 12.6 56.2 37.9 68-10 4 days 26.2 13.0 46.4 32.7

4 Range 7 7-7- / 30 120-105- 90-60- 45-15- 0 30 60 90 120 150 180 210 Days of Age Figure 18. Heights of six whooping cranes

35 TABLE II. LOGISTIC GROWTH CURVE VALUES AND MEANS FOR WEIGHTS OF SIX WHOOPING CRANES Crane No. K Yo A b 68-2 6800 gm. 132 gm. 50.5.528 68-3 5700 136 40,9.501 68-4 6800 118 56.6.543 68-6 5775 108 52.4.533 68-7 6200 131 46.3.517 68-8 6700 133 49.3.525 Mean 6329 126 49.3.525

0000 Y Q800 1 + 50.5 exp.528 t o 0 o 0 0 00 00 1 2 10 11 12 13 14 15 N D-3 (D = Day of Age) Figure 19. Logistic growth curve for crane 68-2

8000 6000 0 0 0 0 00 00 0 4000 0 2000 Y 5700 1 + 40.9 exp -.501 t 000 00 0 o 0 2 3 6 9 110 11 12 13 114 15 D-3 (D = Day of Age) Figure 20. Logistic growth curve for crane 68-3

8000 6000 4000 6800 Y = 1 + 56.6 exp -.543 t 00 0 2 3 4 5 6 7 8 9 10 11 12 13 14 15 N D-3 (D = Day of Age) Figure 21. Logistic growth curve for crane 68-4

4000 Y 5775 = 1 + 52.4 exp -.533 t 2000 o 0 o 000 o 00 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 N D-3 (D = Day of Age) Figure 22. Logistic growth curve for crane 68-6

8000 6000 0 0 0 0 0 0 4000 Y = 6200 1 + 46.3 exp -.517 t 2000 10 11 12 13 14 15 D-3 (D = Day of Age) Figure 23. Logistic growth curve for crane 68-7.N. 0

8000 o 00 0 6000 E 4000 0 o 0 0 0 0 UJ 0 0 Y = 6700 1 + 49.3 exp -.525 t 2000 o 00000 000 0 0 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 N D-3 (D = Day of Age) Figure 24. Logistic growth curve for crane 68-8

8000 6000 E ro c.d 4000 4_1 CU Y = 6329 1 + 49.3 exp -.525 t 2000 10 11 12 13 14 15 N D-3 (D = Day of Age) Figure 25. Mean logistic growth curve for six whooping cranes

43 DISCUSSION The Downy Stage The downy stage of the young cranes lasts about six weeks. The overall down becomes shabbier and grayer and the underparts darken as the birds grow. This change may be due to a bleaching effect by the sunlight, plumage wear, progressive growth of the down with a lighter color appearing at the base, or a combination of these factors. The down lengthens as the bird grows, lengthening at a more rapid rate on the trailing edges of the wings, tail, and scapular area. A long, downy ear tuft becomes noticeable at about four weeks. At 40 days the black primaries are just beginning to appear as pinfeathers. Development of the Juvenal Plumage A dramatic change takes place between the sixth and seventh week when, within the span of a week, the birds attain at least half of their contour feathers. At this point the wings are burdened with the emerging secondaries and primaries and are drooping much of the time. As the juvenal plumage is acquired, the down is not molted but continues to grow longer with the downy portions attached to the tips of the contour feathers. These tips of down eventually break off and the birds are in their full juvenal plumage by 100 days.

44 The Juvenal Molt The time at which the birds were first able to fly was not determined since they were confined in a relatively small area and were either pinioned at an early age or wing clipped shortly after the primaries began to emerge. It is assumed, however, that flight is possible by 100 days of age, at which time the birds are in their full juvenal plumage. There is a gradual molt to the adult, or all white, plumage commencing shortly after the juvenal plumage is attained. Replacement of feathers was first noted on the upper back and ventral portions of the neck at four months. The molt progresses upward on the neck and is complete on the neck and head by 13 months. The secondaries, marginal coverts, part of the primary coverts, underlining of the wings, and feathers of the lower back and rump are initially white on the juvenal and replacement of feathers in these areas was not determined. Feathers of the legs and abdomen were initially tinged with cinnamon but gradually turned white, presumably from wearing away of the outer downy portions. The cinnamon tipped wing coverts began to molt on most of the birds shortly before 300 days and were not completely replaced until late in August (450 days). All of the 1968 hatch birds had replaced all of their wing coverts by this time, but all of the 1967 hatch birds retained a few cinnamon tipped coverts well past August 1968.

The black, hairlike feathers of the head first appear at the base of the mandibles at about 50 days of age, but only become markedly 45 noticeable at 200-250 days. The reddish skin on the head first becomes apparent at about 175 days. The legs and feet are initially very dark, but have lightened considerably by 50 days. After this they gradually darken to the blackish-slate of the adults. The egg tooth is usually lost between two and five weeks. The flesh color (or seashell pink when it first appears) on the proximal portion of bill is first evident at about 40 days. The eyes are dark olive when the chicks first hatch, gradually lightening to massicot yellow in the adults. Height The measurements of height used in figure 18 are estimates and were dependent upon the posture of the bird. All measurements were made as the bird stood in a normal upright position as illustrated in the line drawings (figures one through nine). Although the individual measurements are only approximations, they serve to illustrate growth in cranes. The range of variation in height may be an indication of sex, genetic variation or the effect of disease. (Personal observations indicate a height difference between the sexes in sandhill cranes.)

46 Weight An actual weight gain was not observed until about the third day of age due to the refusal of the birds to eat unless fed by hand. Thus D-3 was used as a basis for the time scale. The transformation N D-3 produced a logistic curve which approximated the growth data, although the curve tends to exceed the actual data during the first month of growth and is slightly lower for each individual towards the end of the growth period. These differences can be expected, however, since this equation is merely a theoretical approximation of a complex set of growth parameters. A dip occurs in the actual weight gain for the six birds from 30 to 60 days of age. This leveling off or decline in weight was the result of infections of coccidia (Eimeria sp..) and an unidentified protozoan which were arrested through medication. Later, minor undulations in weight may have been the result of heat stress during several late summer hot spells. Based on the data, five of the six cranes fall into two weight categories. (Crane 68-7 was intermediate between the two groups.) There is approximately 1,000 grams difference in the final weights of these two groups, which may be an indication of sex, since in some species of cranes the adult males are larger than the adult females (Walkinshaw 1949). Sex differences are not readily apparent in whooping cranes and no attempt was made to sex the birds during the course of the study. The size difference could also be indicative of

47 genetic variation, or retardation of growth caused by a more severe disease infection in some individuals than in others. Table III correlates age, height, weight, and plumage development to facilitate comparison of age classes. The environment created under the captive conditions may have accelerated or inhibited the rate of development compared to the rate in the wild. The study birds most likely experienced greater extremes of temperature since birds in the wild are able to modify the temperature ranges under which they exist by their habits of migration. The diet (a commercial poultry diet) also may have had an influence on the rate of development. Such factors serve to demonstrate the need of further observations on wild birds to help validate this study. The key to age identification is included as an aid in identifying age classes in young whooping cranes under field conditions.

48 TABLE III. PROGRESSIVE CHANGES IN APPEARANCE USEFUL IN IDENTIFYING AGE CLASSES Height Weight Age in cm. in gm. Stage of Development Newly hatched 19.0-20.5 100-150 1 day 19.0-20.5 100-150 5 days 25.5 120-165 11 days 30.5 200-300 18 days 33.0-40.5 350-625 28 days 48-53 850-1,050 40 days 65-75 1,750-2,500 50 days 80-98 2,450-3,350 Down wet. Legs and feet swollen, reddish. Whitish egg tooth present on tip of upper mandible. Down dry. Swelling gone from legs and feet, dark in appearance. Eye dark olive. (Figure 1, Figure 10) Eye olivaceous black with a glassy bluish cast. (Figure 2) Egg tooth still present. (Figure 3) "V" of pale pinkish cinnamon on breast. Egg tooth may or may not be present. (Figure 4) Downy ear tuft very noticeable, one-fourth inch in length. Eye olive. Down of scapular area, trailing edge of wing, and tail longer than other areas. Legs lighter in appearance. Egg tooth gone. (Figure 5) Black primaries appearing as pinfeathers. Still entirely downy. (Figure 6, Figure 10) Plumage half contour feathers and half down. Black hairlike feathers are present at the base of the mandibles. (Figure 7, Figure 11)

49 Age Height in cm. Weight in gm. Stage of Development 75 days 100-115 3,950-4,650 100 days 100-120 4,300-4,950 125 days 4,825-5,700 150 days 5,200-6,075 175 days 5,500-6,400 200 days 112-127 5,700-6,800 Contour feathers of the head and neck beginning to emerge. Legs still downy. Retrices nearly three inches long with downy tips. Decurved tertials extend down onto the tail. (Figure 8, Figure 11, Figure 14) Complete covering of contour feathers. Full juvenal plumage. A light scattering of black hairlike feathers is visible on the cheek beneath the eye. (Figure 9, Figure 12, Figure 14) Black on forehead as well as cheek. Eye olive-buff. A few feathers have been renewed on the upper back and ventral portion of the neck. Legs and feet blackish-slate, as in the adults. (Figure 17) Black much more pronounced on cheek and forehead. Half of juvenal feathers replaced on upper back of most birds. Increase in replacement of feathers on lower half of neck and breast. (Figure 12) Carrot red skin shows through black on forehead. Eye cream color. Replacement of feathers on upper neck gives a salt and pepper effect. Black now visible on crown. Some of head plumage replaced giving a white flecked appearance. Upper back has only a scattering of cinnamon feathers remaining. (Figure 15)

50 Aaa Height in cm. Weight in gm. Stage of Development 250 days 112-127 300 days 112-127 350 days 112-127 400 days 112-127 450 days 112-127 Large whitish patch evident behind and below the eye, bordering on the black cheek patch. (Figure 15) A scattering of glossy black feathers (not the hairlike feathers) is present on the rear portion of the crown and back of the head. Eye massicot yellow. Back completely white. From none to three-fourths of the middle wing coverts have been replaced. (Figure 13) A scattering of cinnamon persists on the upper neck and head. Only a few cinnamon wing coverts remain. (Figure 16) A few cinnamon wing coverts still remain. Head and neck entirely white. Plumage as in the adults. Entirely white, no cinnamon remaining. (Figure 13, Figure 16)

51 KEY TO THE AGE OF YOUNG WHOOPING CRANES 1. Plumage all downy 2 Plumage not all downy; contour feathers present 6 2. Plumage wet; legs and feet swollen and reddish... NEWLY HATCHED Plumage dry; legs and feet not swollen and reddish 3 3. Less than 26 centimeters tall ONE TO FIVE DAYS More than 26 centimeters tall 4 4. Downy ear tuft not long and noticeable; egg tooth may or may not be present SEVEN TO TWENTY-ONE DAYS Downy ear tuft long and noticeable; egg tooth not present 5 5. Black primaries not yet visible; down of scapular area, tail, and trailing edge of wing longer than other areas.... THIRTY DAYS Black primaries visible as pinfeathers FORTY-TWO DAYS 6. Plumage one half contour feathers and one half down (see Figure 11) FIFTY DAYS Plumage more than one half contour feathers....... 7 7. Legs still downy; contour feathers of head and neck beginning to emerge SEVENTY-FIVE DAYS No down remaining; entire plumage composed of contour feathers 8 8. No replacement of juvenal plumage; black visible on cheek only (full juvenal plumage) ONE HUNDRED DAYS Some or all of juvenal plumage replaced by white plumage; black visible on forehead as well as cheek 9 9. Middle wing coverts not replaced; back not entirely white. 10 Middle wing coverts at least partially replaced; back entirely white 13 10. Less than one half of feathers replaced on upper back and ventral portion of neck (see Figure 17).. ONE HUNDRED TWENTY-FIVE DAYS One half or more of feathers replaced on upper back (see Figure 12) 11 11. No plumage replacement on head; black more pronounced on cheek and forehead, but not visible on crown.. ONE HUNDRED FIFTY DAYS Plumage replacement on head; black visible on crown 12 12. Large whitish patch behind eye not yet distinct; white flecked appearance on head (see Figure 15) TWO HUNDRED DAYS Large whitish patch evident behind and below eye, bordering on black cheek patch (see Figure 15) TWO HUNDRED FIFTY DAYS

52 13. Less than three-fourths of middle wing coverts replaced* (see drawing below) THREE HUNDRED DAYS More than three-fourths of middle wing coverts replaced... 14 14. Head and neck having scattered cinnamon feathers THREE HUNDRED FIFTY DAYS Head and neck entirely white 15 15. Plumage not all white; a few cinnamon wing coverts remain... FOUR HUNDRED DAYS Plumage all white; adult plumage FOUR HUNDRED FIFTY DAYS AND OLDER *One individual had not molted any middle wing coverts by this time.

53 LITERATURE CITED Allen, R. P. 1952. The whooping crane. Research Rept. 3, Natl. Audubon Soc. New York. 246 pp. Bent, A. C. 1926. Life histories of North American marsh birds. U. S. Natl. Mus. Bull. 138. Washington, D. C. 490 pp. Biester, H. E., and L. H. Schwarte. 1959. Diseases of poultry. Iowa State Univ. Press, Ames. 1103 pp. Blyth, E., and W. B. Tegetmeier. 1881. The natural history of the cranes. Cox and Porter, London. 91 pp. Kendall, M. G., and W. R. Buckland. 1966. A dictionary of statistical terms. 2nd ed. Oliver and Boyd. Edinburgh, London. 575 pp. Ridgway, R. 1912. Color standards and color nomenclature. R. Ridgway, Washington, D. C. 43 pp. Ridgway, R., and H. Friedmann. 1941. The birds of Middle and North America. U. S. Natl. Mus. Bull. 50. Washington, D. C. 245 pp. Stevenson, J. 0., and R. E. Griffith. 1946. Winter life of the whooping crane. Condor 48(4): 160-178. Walkinshaw, L. H. 1949. The sandhill cranes. Cranbrook Inst. Sci. Bull. 29. Bloomfield Hills, Michigan. 202 pp.

APPENDIX I

54 Colors Used in the Text From Robert Ridgway's Color Standards and Color Nomenclature COLOR PLATE NUMBER* benzo brown blackish slate bone brown carnelian red carrot red chaetura black chaetura drab cinnamon cinnamon - brown XLVI LII XL XIV XIV XLVI XLVI XXXI XV cinnamon buff XXIX clay color clove brown cream color dark grayish olive dark olive XXIX XL XVI XLVI XL dark olive gray LI deep grayish olive drab XLVI XLVI dull purplish black dull violet - black flesh color XLIV XIV *from Ridgway

55 grayish olive grenadine pink COLOR PLATE NUMBER XLVI II light cinnamon drab XLVI light drab light grayish olive light grayish vinaceous light ochraceous - buff XLVI XLVI XXXIX XV light ochraceous salmon XV light olive - gray light pinkish cinnamon light purplish vinaceous light salmon - orange light vinaceous - cinnamon light vinaceous - fawn light yellowish olive massicot yellow mummy brown olivaceous black (1) olive olive - buff orange - pink pale flesh color pale ochraceous - buff pale pinkish buff LI XXIX XXXIX II XXIX XL XXX XVI XV XLVI XXX XL II XIV XV XXIX

56 COLOR pale pinkish cinnamon pale salmon color PLATE NUMBER XXIX XIV peach red pinkish buff pinkish cinnamon salmon - buff sayal brown seashell pink sepia smoke gray XXIX XXIX XIV XXIX XIV XXIX XLVI vinaceous fawn XL yellowish olive XXX