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OF the RING-NECKED PHEASANT DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosoph

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OF the RING-NECKED PHEASANT DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosoph observations on the renesting behavior OF THE RING-NECKED PHEASANT DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By John Lyman Seubert, B. S., M. S. The Ohio State University 1956 Approved by: Adviser Department of Zoology and Entomology Acknowledgements The author is deeply indebted to Dr. Eugene H. Dustman, Leader, the Ohio Cooperative Wildlife Research Unit, for his generous and able guidance and constant interest during this study. The interest and valuable counsel of Dr. Charles A. Dambach, Director of Natural Resources Institute, The Ohio State University, has been greatly appreciated. Special thanks are due to Mr. Walter Hoy, statistician, the Statistics Laboratory, The Ohio State University, for the statistical analyses used in this manuscript. I am indeed grateful to Mr. Henry Marquette, owner of the farm property upon which the research area was located, for his interest and cooperation. To the many persons who offered their advice and assistance in the field, I extend my thanks, particularly to Dr. Ernest E. Good, Eugene Knoder, James Hume, and Ralph Andrews. I am grateful for the financial assistance in the form of a Research Fellowship granted by the Ohio Cooperative Wildlife Research Unit and for the undeiwriting of the costs of this study by this agency. To my wife, Jean, my heartfelt thanks for her encouragement, patience, helpful suggestions, and for the typing and proofreading of this manuscript. Table of Contents Section Acknowledgements ....... ......... ........ H X. Introduction ....... ...... ..... l A. Perspective......... 1 5. Objectives ......... ........ ....... 5 II. Description of the Study Area and Methods of Investigation ......................... 7 A. Location of Study Area ......... ....... 7 1. Physiographic and Geologic Setting ....... 7 2. Soils ............. ...... 9 3. Climate ........ 9 B. Description of the Experimental Enclosure .... 9 1. Size and Fencing .......o...... 9 2. Vegetation on the Study Area ........ 11 a. Type and Arrangement of Vegetation . 11 b. Control of Vegetative Growth ...... 11 3. Predator Control Measures .......... 13 C. Study Methods ..... ............ ..... 13 1. Brood Stock 13 2. Observations of Brood Stock ......... II4. 3. Location of Nests ....... .......... 17 I4.. Aging of Clutches and the Assignment of Nests to Categories of Incubation......... 17 5. Disruption of Hens ............. 20 iii iv Section III. Results . ......... ............... 23 A. Behavior of Brood Stock in Relation to the Enclosure A r e a .......................... 23 B. Breeding Behavior ........................... 23 1. Territorialism .................... 2k a. Territorial Conflict ........... 2$ b. Aggressiveness and Dominance......... 25 c. Territorial Establishment ....... 27 1. Time of Establishment........... 27 2. Number and Sizes of Territories • . 29 3. Fluctuations in the Sizes of Territories.................. 29 U. Territorial Requirements ....... 29 5. Cessation of Territorialism .... 32 2. Non-Territorial Cocks ..... ........... 33 3. Harem Formation ........................ 35 a. Number and Sizes of Harems ....... 35 b. Stability of Harem Composition ..... 36 c. Dominance Hierarchy of Hens ...... 36 I4. Territory and Harem Relationships......... 39 5. Inter-Specific Strife .............. J4O 0. Egg Laying, Nesting, and Incubation........... 1*0 1. Hens Available for Nesting ......... Itl 2. Nest Establishment and Incubation ...... U6 a. Number of Nests Per Hen U6 V Section b. Nest Types and Clutch Sizes ...... U8 3* Egg Production and Fertility ....... Time and Duration of the Nesting Period • • . 55 D. Behavior of Nesting Hens , ............. 57 1. Behavior of Incubating Hens ......... 57 2. Behavior of Disrupted Hens ......... 60 3* Clutch Usurpation and Adoption ......... 62 U* Nest Desertion and Abandonment ....... 62 5- Location of Nests in Territories and in Cover Stripe ..................... 6I4. 6 . Renesting .................. 67 a. Nest Re-Establishment in Relation to Stage of Incubation .......... 67 b. Renesting Interval ...... ........ 6? c. Avian Ovulation ......... 75 d. Renesting in Relation to Time of Season. 82 e. Reproductive Success in Relation to Renesting and Hatching Peaks ......... 90 f. Renesting Activity as Indicated by Nest Establishment and Hatching Curves , . 101 E. Behavior of Hens with Broods ............ 106 1. Number of Clutches H a t c h e d ........... 106 2. Behavior of Hens at Hatching ........... 106 3* Movements of Broods 107 1|. Brood Defense .................. 107 vi Section $. Adoption of Broods ............. 107 F. Bird L o s s e s ................................. 107 IV. Summary and Conclusions ................. 112 V. Literature Cited .............. ........ 119 Autobiography............ 127 List of Tables Table 1. Dates of Estaolishment and. Sizes of Territories and Dates of Establishment and Sizes of Harems, 1950-1951....................................... 28 2. Harem Composition and Degree, of Initial Nesting, 1950-1951 ............................ 37 3- Nesting Data, 1950-1951 1+5' U. Nest Establishment and Incubation, All Nests, 1950-1951 .......................................... 1+7 5. Mean Number of Eggs for Various Types of Nests, 1950-1951 ......................................... 51 6. Clutch Sizes of First Nests and Renests, 1950-1951 • • • 53 7« Egg Production, 1950-1951 ............................ 55 8. Nest Establishment Data, All Nests, 1950-1951 58 9. Nest Desertion and Abandonment, 1950-1951 . ....... 83 10. Nesting of Harem Hens in Relation to Territories, 1950-1951 ...... ............................... 65 11. Hens Renesting After Disruption From Initial and Second Nests, 1950-1951 68 12. Hens Renesting After Disruption From Initial and Second Nests, 1950-1951 Data Combined ................ 70 13* Hens Renesting After Disruption or Desertion, 1950 . 83 ll+» Hens Renesting After Disruption or Desertion, 1951 . 81+ vii viii Table 15. Pheasant Productivity as Affected by Renesting, Hatching Chronology, and Nest Disruption at Various Times During the Nesting Season ..... ........... 9U 16. Dates of Initial Nest Establishment, 1950-1951....... 95 17. Potential Hatching Distribution, 1950-1951 ........... 96 18. Adult Bird Losses, 1950-1951 ........................ 108 List of Figures Figure 1. Map of Ohio Showing General Location of Research Ar e a ................................... 8 2. The Research A r e a ................................ 10 3* View of the Research Area Looking Northeast......... j.2 J+* Attaching Plastic Marker to Hen Before Release....... l£ 5. Hen with Plastic Markers and Leg Bands ....... 16 6. Incubating Hen .................................. 18 7. Aging E m b r y o s .................................... 19 8 . Hen Disruption Procedure ..................... 21 9. Site of Disrupted N e s t ............................. 22 10. Cocks Fighting .......... .................. 26 11. Cock Territories, 1 9 5 0 ............................. 30 12. Cock Territories, 1 9 5 1 ............................. 31 13. Territorial Cock and Part of H ar em.................. 38 1U. Dropped Eggs ............. U2 15. Dump Nest in Clump of Red Clover................... I43 16. Incubated N e s t .................................... UU 17. Compound Dump Nest , .......... ip? 18. Periods of Nest Establishment, 1950-1951 ............ 56 19* Display of Pugnacious H e n .......... 61 20. Regression of Length of Renesting Interval on Stage of Incubation at Destruction of First Nest, 1950-1951 . lb 21. Renesting in Relation to Time of Season ....... 86 ix X Figure 22. Nest Estaolishment Data, Buss (19U6)................ 102 23. Nest Establishment Data, Initial Nests, 19$0-19$! . 10U 2k* Nest Establishment Data, All Nests, 1950-19$1....... 105 25. Avian K i l l ...................................... Ill I. Introduction A. Perspective Renesting following unsuccessful nesting attempts has been accepted as a common occurrence in the nesting behavior of the ring­ necked pheasant (Phasianus colchicus torquatus) which typically raises one brood each year. Although there are many references in the liter­ ature about the occurrence or probable occurrence of renesting by pheasants as well as by other avian species, few are based upon direct observation. Most of the investigators have inferred renesting on the basis of indirect evidence, i.e., final reproductive success, clutch sizes, time of season, type of nest site, and the amount of down present in nests (duck nests). Hamerstrom (1936) considered some of the pheasant nests found in small grain as probable renesting attempts because of the lateness of the season. Errington and Hamerstrom (1937) considered the theoretical aspects of renesting, and evaluated nesting losses of pheasants in relation to brood success. They suggested that by renestings, a great majority of hens succeed in bringing off noim- ally satisfactory broods in spite of heavy nesting losses. Randall (I9I4O) apparently distinguished renests by smaller’ clutch size and -1- time of season and wrote that by renesting efforts, between 52 and 55 percent of the hen pheasants on his study area raised broods, although only 25 percent of all nests were successful. Knott et al. (19U3 ) reported that a study of pheasant brood records for the year 19U0 snowed that eight percent of the broods resulted from renesting attempts, these broods averaging 3*8 young each as compared to 8.2 young from first attempts.
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