Status of the Red Knot (Calidris Canutus Rufa) in the Western Hemisphere
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Status of the Red Knot ( STATUS OF THE RED KNOT (CALIDRIS CANUTUS RUFA) IN THE WESTERN HEMISPHERE Calidris canutus rufa LAWRENCE J. NILES, HUMPHREY P. SITTERS, AMANDA D. DEY, PHILIP W. ATKINSON, ALLAN J. BAKER, KAREN A. BENNETT, ROBERTO CARMONA, KATHLEEN E. CLARK, NIGEL A. CLARK, CARMEN ESPOZ, PATRICIA M. GONZÁLEZ, BRIAN A. HARRINGTON, DANIEL E. HERNÁNDEZ, KEVIN S. KALASZ, RICHARD G. LATHROP, RICARDO N. MATUS, CLIVE D. T. MINTON, R. I. GUY MORRISON, ) Niles et al. Studies in Avian Biology No. 36 MARK K. PECK, WILLIAM PITTS, ROBERT A. ROBINSON, AND INÊS L. SERRANO Studies in Avian Biology No. 36 A Publication of the Cooper Ornithological Society STATUS OF THE RED KNOT (CALIDRIS CANUTUS RUFA) IN THE WESTERN HEMISPHERE Lawrence J. Niles, Humphrey P. Sitters, Amanda D. Dey, Philip W. Atkinson, Allan J. Baker, Karen A. Bennett, Roberto Carmona, Kathleen E. Clark, Nigel A. Clark, Carmen Espoz, Patricia M. González, Brian A. Harrington, Daniel E. Hernández, Kevin S. Kalasz, Richard G. Lathrop, Ricardo N. Matus, Clive D. T. Minton, R. I. Guy Morrison, Mark K. Peck, William Pitts, Robert A. Robinson, and Inês L. Serrano Studies in Avian Biology No. 36 A PUBLICATION OF THE COOPER ORNITHOLOGICAL SOCIETY Front cover photograph of Red Knots by Irene Hernandez Rear cover photograph of Red Knot by Lawrence J. Niles STUDIES IN AVIAN BIOLOGY Edited by Carl D. Marti 1310 East Jefferson Street Boise, ID 83712 Spanish translation by Carmen Espoz Studies in Avian Biology is a series of works too long for The Condor, published at irregular intervals by the Cooper Ornithological Society. Manuscripts for consideration should be submitted to the editor. Style and format should follow those of previous issues. Price $20.00. To order, add $2.75 for postage in the US, $9.75 for Canada, and $11.75 for every- where else. Make checks payable to the Cooper Ornithological Society (in U.S. funds), or pay by MasterCard or Visa. Send orders to Cooper Ornithological Society, c/o John Rotenberry, Department of Biology, University of California, Riverside, CA 92521 USA. Credit card orders may be sent via email to: [email protected]. Permission to Copy The Cooper Ornithological Society hereby grants permission to copy chapters (in whole or in part) appearing in Studies in Avian Biology for personal use, or educational use within one’s home institution, without payment, provided that the copied material bears the statement “©2008 The Cooper Ornithological Society” and the full citation, including names of all authors. Authors may post copies of their chapters on their personal or institutional website, except that whole issues of Studies in Avian Biology may not be posted on websites. Any use not specifi cally granted here, and any use of Studies in Avian Biology articles or portions thereof for advertising, republication, or commercial uses, requires prior consent from the editor. ISBN: 978-0-943610-83-2 Library of Congress Control Number: 2008929213 Printed at Cadmus Professional Communications, Ephrata, Pennsylvania 17522 Issued: 15 June 2008 Copyright © by the Cooper Ornithological Society 2008 ii CONTENTS AUTHOR ADDRESSES ...................................................................................................... xvii ABSTRACT .......................................................................................................................... 1 TAXONOMY ....................................................................................................................... 5 GENETIC EVIDENCE FOR KNOT SUBSPECIES ........................................................................... 7 ECOLOGICAL AND MORPHOLOGICAL EVIDENCE FOR SUBSPECIES IN NORTH AMERICA ............. 9 EVIDENCE FOR DISCRETE WINTERING POPULATIONS .............................................................. 9 PHYSICAL DESCRIPTION ............................................................................................... 11 FIRST BASIC PLUMAGE ......................................................................................................... 11 DEFINITIVE ALTERNATE PLUMAGE ........................................................................................ 13 ALTERNATE PLUMAGE ......................................................................................................... 13 FEMALES ............................................................................................................................. 13 MALES—CALIDRIS CANUTUS RUFA ........................................................................................ 13 MALES—CALIDRIS CANUTUS CANUTUS .................................................................................. 13 MALES—CALIDRIS CANUTUS ROGERSI ................................................................................... 13 MALES—CALIDRIS CANUTUS ISLANDICA ................................................................................ 14 MALES—CALIDRIS CANUTUS ROSELAARI ................................................................................ 14 FIRST ALTERNATE PLUMAGE ................................................................................................ 14 JUVENILE PLUMAGE ............................................................................................................. 14 HATCHLINGS ....................................................................................................................... 14 DISTRIBUTION IN TIME AND SPACE .......................................................................... 14 THE ANNUAL CYCLE ........................................................................................................... 14 BREEDING RANGE ................................................................................................................ 15 WINTER (NON-BREEDING) RANGE ........................................................................................ 17 MIGRATORY RANGE AND MAJOR STOPOVER AREAS .............................................................. 19 BIOLOGY AND NATURAL HISTORY ........................................................................... 25 REPRODUCTION ................................................................................................................... 25 Mating displays ............................................................................................................. 26 Sexual behavior .............................................................................................................. 26 Nest sites ....................................................................................................................... 27 iii Number of broods .......................................................................................................... 27 Clutch size ..................................................................................................................... 27 Incubation period ........................................................................................................... 27 Nestling period .............................................................................................................. 27 Nesting success .............................................................................................................. 28 MORTALITY ......................................................................................................................... 28 LONGEVITY .......................................................................................................................... 28 SITE FIDELITY ...................................................................................................................... 28 MIGRATION ......................................................................................................................... 28 FEEDING HABITS .................................................................................................................. 29 HABITAT ............................................................................................................................. 30 PREFERRED MICROHABITATS ................................................................................................. 30 BREEDING HABITAT ............................................................................................................. 30 MIGRATION AND STOPOVER HABITAT—CANADA ................................................................. 30 MIGRATION AND STOPOVER HABITAT—NORTHEASTERN U.S. ............................................... 30 MIGRATION AND STOPOVER HABITAT—DELAWARE BAY, U.S. ............................................. 30 Habitats important for Red Knots in Delaware Bay ...................................................... 33 Red Knot feeding ecology in Delaware Bay ................................................................... 36 Mapping horseshoe crab spawning habitat suitability ................................................... 39 Mapping critical Red Knot habitat ................................................................................ 41 Evidence of decline in both the population of horseshoe crabs and the availability of their eggs for Red Knots ......................................................................................... 44 MIGRATION AND STOPOVER HABITAT—SOUTHEASTERN UNITED STATES ............................... 47 MIGRATION AND STOPOVER HABITAT—PANAMA ................................................................. 48 MIGRATION AND STOPOVER HABITAT—MARANHÃO, BRAZIL ..............................................