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Temperature, Nest Humidity, and Behavioral Thermoregulation in a Hot Environment

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Temperature, Nest Humidity, and Behavioral Thermoregulation in a Hot Environment (ISBN: 0-943610-30-3) AVIAN INCUBATION: EGG TEMPERATURE, NEST HUMIDITY, AND BEHAVIORAL THERMOREGULATION IN A HOT ENVIRONMENT BY GILBERT S. GRANT Departmentof Biology,University of California, Los Angeles ORNITHOLOGICAL MONOGRAPHS NO. 30 PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION WASHINGTON, D.C. 1982 AVIAN INCUBATION: EGG TEMPERATURE, NEST HUMIDITY, AND BEHAVIORAL THERMOREGULATION IN A HOT ENVIRONMENT ORNITHOLOGICAL MONOGRAPHS This series, publishedby the American Ornithologists'Union, has been estab- lished for major paperstoo long for inclusionin the Union's journal, The Auk. Publicationhas been made possible through the generosity of thelate Mrs. Carll Tucker and the Marcia Brady Tucker Foundation, Inc. Correspondenceconcerning manuscriptsfor publication in the series should be addressedto the Editor, Dr. Mercedes S. Foster, USFWS, NHB-378, National Museum of Natural History, Washington, D.C. 20560. Copies of Ornithological Monographs may be ordered from the Assistant to the Treasurer of the AOU, Glen E. Woolfenden, Department of Biology, Uni- versity of South Florida, Tampa, Florida 33620. (See price list on back and in- side back cover.) Ornithological Monographs, No. 30, ix + 75 pp. Editor of AOU Monographs, Mercedes S. Foster Special Reviewers for this issue, Cynthia Carey, Department of Environ- mental, Population, and Organismic Biology, University of Colorado, Boulder, Colorado; Donald F. Hoyt, Department of Biological Sci- ences,California State PolytechnicUniversity, Pomona, California; S. Charles Kendeigh, Department of Ecology, Ethology, and Evolu- tion, University of Illinois, Champaign, Illinois Author, Gilbert S. Grant; present address: North Carolina State Museum of Natural History, P.O. Box 27647, Raleigh, North Carolina 27611 First received, 11 August, 1981; accepted, 29 December, 1981; final re- vision completed, 9 June, 1982 Issued 2 November 1982 Price $9.00 prepaid ($7.00 to AOU members) Library of Congress Catalogue Card Number 82-73590 Printed by the Allen Press, Inc., Lawrence, Kansas 66044 Copyright ¸ by the American Ornithologists'Union, 1982 ISBN: 0-943610-30-3 AVIAN INCUBATION: EGG TEMPERATURE, NEST HUMIDITY, AND BEHAVIORAL THERMOREGULATION IN A HOT ENVIRONMENT BY GILBERT S. GRANT Department of Biology, University of California, Los Angeles ORNITHOLOGICAL MONOGRAPHS NO. 30 PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION WASHINGTON, D.C. 1982 TABLE OF CONTENTS INTRODUCTION .................................................................................. 1 STUDY AREA ...................................................................................... 1 AMBIENT WEATHER DATA ................................................................ 2 SALINITY .......................................................................................... 3 MATERIALS AND METHODS .............................................................. 4 EGG AND NEST DATA ...................................................................... 4 TEMPERATURE DATA ........................................................................ 6 HUMIDITY DATA .............................................................................. 7 SOLAR RADIATION ............................................................................ 8 WIND .............................................................................................. 8 WATER UPTAKE OF FEATHERS .......................................................... 8 OXYGEN CONSUMPTION OF EGGS ...................................................... 8 EGG REFLECTANCE .......................................................................... 9 TEMPERATURE OF COPPER STILT CHICK ............................................ 9 TIMING OF BREEDING ........................................................................ 9 THERMAL BIOLOGY OF THE EGG .................................................... 11 NEST AND EGG PARAMETERS ............................................................ 11 NEST AIR AND EGG TEMPERATURES .................................................. 13 EGG HEATING EXPERIMENTS ............................................................ 15 LETHAL EGG TEMPERATURES ............................................................ 17 CLUTCH SIZE AND EGG TEMPERATURE .............................................. 18 EGG WATER LOSS AND NEST HUMIDITY ........................................ 21 PHYSICAL DIMENSIONS OF EGGS AND SHELLS .................................... 21 INCUBATION PERIOD ........................................................................ 22 EGG MASS LOSS AND WATER-VAPOR CONDUCTANCE .......................... 23 FRACTIONAL MASS LOSS .................................................................. 25 EGG, NEST, AND AMBIENT VAPOR PREssuRE .................................... 25 NEST VENTILATION .......................................................................... 26 EMBRYONIC OXYGEN CONSUMPTION ............................................ 28 EFFECTS OF SALT AND MUD ON EGGS .......................................... 29 INCUBATION BEHAVIOR .................................................................. 33 EGG-COVERING BEHAVIOR ................................................................ 33 BELLY-SOAKING AND FOOT-WETTING ................................................ 35 ORIENTATION ON THE NEST .............................................................. 40 PANTING AND GULAR FLUTTERING .................................................. 43 PTERYLOEKECTION ............................................................................ 43 ATTENTIVE BEHAVIOR ...................................................................... 45 TEMPERATURE OF AN INCUBATING STILT .......................................... 53 EFFECT OF BEHAVIOR ON EGG TEMPERATURE .................................. 54 USE OF SHADE .................................................................................. 57 STILT CHICK THERMOREGULATION .............................................. 59 DISCUSSION ........................................................................................ 59 THERMOREGULATION OF EGGS, CHICKS, AND ADULTS ...................... 60 REGULATION OF NEST HUMIDITY .................................................... 62 FUNCTION AND ORIGIN OF BELLY-SOAKING ...................................... 65 ACKNOWLEDGMENTS ........................................................................ 69 SUMMARY ............................................................................................ 69 LITERATURE CITED ............................................................................ 70 vi LIST OF FIGURES Figure 1. Map of southernCalifornia showinglocations of study areas and weather stations ...................................................................... 2 2. Monthly mean ambient temperatures, relative humidities, and rainfall near the Salton Sea ...................................................... 3 3. Shaded ambient temperatures and relative humidities at ground level near nestsat the Salton Sea at different times of day ........ 4 4. Ambient temperatures and relative humidities near the coastal study sites................................................................................ 5 5. Nesting chronology of the Killdeer at the Salton Sea ................ 10 6. Nesting chronologyof the Black-necked Stilt at the Salton Sea .... 11 7. The relationshipbetween ambient temperatureand incubatedegg temperature of the Killdeer ...................................................... 15 8. The relationshipbetween ambient temperature and incubated egg temperature of the Lesser Nighthawk ...................................... 16 9. The relationshipsbetween ambient temperature and incubated egg temperatureof the American Avocet at two sites................ 17 10. The relationshipsbetween ambient temperature and incubated egg temperature of the Forster's Tern nesting in two habitats __ 18 11. Relationship between ambient temperature and incubated egg temperature of the Black-necked Stilt ...................................... 19 12. Temperatures of a pipped egg of the Black-necked Stilt ............ 21 13. The relationship between oxygen consumptionby eggs of the Black-necked Stilt and days incubated ...................................... 30 14. Nest of the Black-neckedStilt at the SaltonSea showing three mud-covered eggs and a large clump of mud in the nest ............ 31 15. Temperaturesof chicken eggsexposed to various environmental and experimental conditions at the Salton Sea .......................... 34 16. Mean vapor pressure at ground level near stilt nests and mean number of belly-soaksper nest ................................................ 35 17. Frequency of belly-soaking and temperatures at one Black- necked Stilt nest ...................................................................... 36 18. Black-necked Stilts at nest relief .............................................. 37 19. Frequency of belly-soaking,attentiveness, and temperaturesat an American Avocet nest at the Salton Sea .............................. 38 20. Meteorological conditions, egg temperatures, and behavior at two Forster's Tern nests .......................................................... 39 21. Meteorological conditions, egg temperatures, and behavior at a Forster's Tern nest .................................................................. 40 22. Attentive behavior of Snowy Plovers in relation to time of day, ambient temperature, vapor pressure, and relative humidity
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