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UNIVERSITY of CALIFORNIA, SAN DIEGO Phainopepla Nestlings UNIVERSITY OF CALIFORNIA, SAN DIEGO Phainopepla nestlings adjust begging behaviors to different male and female parental provisioning rules A thesis submitted in partial satisfaction of the requirements for the Master of Science degree in Biology by Jeanne Marie Messier Committee in charge: Professor Sandra L. Vehrencamp, Chair Professor David S. Woodruff Professor Joshua R. Kohn Professor Trevor D. Price 2000 The Thesis of Jeanne Marie Messier is approved, and it is acceptable in quality and form for publication on microfilm and electronically: __________________________________________________________ __________________________________________________________ __________________________________________________________ __________________________________________________________ Chair University of California, San Diego 2000 iii DEDICATION This Thesis is dedicated to Anne and John Messier, in memory of their daughter Jeanne. iv TABLE OF CONTENTS Signature page........................................................................................................ iii Dedication.............................................................................................................. iv Table of Contents................................................................................................... v List of Figures........................................................................................................ vi List of Tables......................................................................................................... vii Preface................................................................................................................... viii Acknowledgements............................................................................................... x Vita........................................................................................................................ xi Abstract................................................................................................................. xii Introduction........................................................................................................... 1 Methods................................................................................................................. 5 Results................................................................................................................... 13 Discussion............................................................................................................. 28 References............................................................................................................. 34 v LIST OF FIGURES Figure 1. Provisioning and begging variables in subsequent parental visits............ 10 Figure 2. Feeding rates and weight gain for deprived nestlings and nestmates....... 16 Figure 3. Food allocation to deprived nestling versus its previous weight loss....... 17 Figure 4. Changes in begging behaviors between control and experimental sessions for deprived and non-deprived nestlings in male- and female-tended nests.......................................................................................................................... 23 vi LIST OF TABLES Table 1. Parental provisioning rates in control and experimental sessions.............. 19 Table 2. Results of multiple linear regression analyses on parental allocation of feeds as a function of nestling begging behaviors.............................................. 21 Table 3. Summary of primary nestling begging behaviors and cues used by parents to allocate food in each nest........................................................................ 26 vii PREFACE Jeanne Messier’s plan for her PhD dissertation research was to investigate the honesty of parent-offspring communication system in two avian species: one species with a small clutch of synchronously hatching chicks, which was predicted to exhibit honest signaling of chick hunger and reliable responsiveness by the parents; and another species with a larger clutch of asynchronously hatching chicks, which was predicted to exhibit some competition among the chicks, possible deceitful signaling of hunger, and parental discounting of chick begging signals. She selected the phainopepla and the robin, respectively. These species had open-cup nests for ease of videotaping, sexually dichromatic adults for distinguishing male and female parents, and non-overlapping breeding seasons. In 1993, her first full field with the early-breeding phainopepla, Jeanne succeeded in conducting nestling removal and replacement studies at 8 nests. Back in the lab, she trained an undergraduate assistant to score the nestling and parental behaviors in recorded videos, then headed up to northern California to work on the robins. Toward the end of the season she was stricken by a hantavirus infection, and was airlifted to a hospital in Reno where she fought for her life. Tragically, she lost. As her advisor, I endeavored to complete the analysis of the phainopepla data. The undergraduate student finished the video data extraction, and I ran the statistical analyses that Jeanne and I had planned, wrote a manuscript draft, and submitted it for publication to a behavior journal. Unfortunately, the manuscript was rejected because of the small sample size. The research project clearly would have benefitted from one more field season. Nevertheless, some highly interesting and statistically significant patterns emerged. She would have been among the first to show parental sex differences viii in provisioning responses to nestlings, and furthermore to show that nestlings learn and adapt their begging behaviors to these sex-specific parental characteristics. Given Jeanne’s well-designed study and skill in executing the field experiments, coupled with her outstanding performance on her earlier qualifying exam, the committee agrees that Jeanne Messier is fully deserving of a posthumous Master of Science in Biology degree. The committee verifies the academic integrity of this thesis submission, which is based on the submitted manuscript. ix ACKNOWLEDGEMENTS We thank Erin Muntean for assistance in the field, Danton Kono and Catherine deRivera for assistance with the video analysis, and Professor Robert Schwager (Cornell University) for statistical advice. Professors Arnon Lotem, Jonathan Wright, and Rebecca Kilner provided stimulating discussions, reviews of manuscript drafts, and access to unpublished manuscripts. The field research was supported by the Los Angeles Audubon Society, Sigma Xi, the UC Natural Reserve System, and training grant support to JMM from the UCSD Biology department. x VITA 1988 BA, Biology, Cornell University 1990-1993 PhD Candidate, Biology, UC San Diego 2000 MS, Biology, UC San Diego WORK EXPERIENCE 1988-1990 Research Assistant, Harvard Medical School PUBLICATIONS Shaw LM, Messier JM, Mercurio AM. 1990. The activation dependent adhesion of macrophages to laminin involves cytoskeletal anchoring and phosphorylation of the alpha-6-beta-1-integrin. Journal of Cell Biology 110:2167-2174. Woo HJ, Shaw LM, Messier JM, Mercurio AM. 1990. The major non-integrin laminin binding protein of macrophages is identical to carbohydrate binding protein-35 (MAC-2). Journal of Biological Chemistry 265:7097-7099. Messier JM, Shaw LM, Chafel M, Matsudaira P, Mercurio AM. 1993. Fimbrin localized to an insoluble cytoskeletal fraction is constitutively phosphorylated on its headpiece domain in adherent macrophages. Cell Motility and the Cytoskeleton 25:223-233. Morin PA, Messier JM, Woodruff DS. 1994. DNA extraction, amplification, and direct sequencing from hornbill feathers. Journal of the Science Society of Thailand 20:31-41. FIELDS OF STUDY Behavioral ecology, animal communication, ornithology xi ABSTRACT OF THE THESIS Phainopepla nestlings adjust begging behaviors to different male and female parental provisioning rules by Jeanne Marie Messier Master of Science in Biology University of California, San Diego, 2000 Professor Sandra L. Vehrencamp, Chair We studied the nestling begging behaviors and parental provisioning responses of the phainopepla (Phainopepla nitens), a sexually dichromatic silky flycatcher native to Mexico and the southwestern United States deserts. Because of its small clutch size (usually two eggs) and synchronous hatching strategy, we predicted that size-based nestling competitive interactions and brood reduction strategies would be absent, so that parental food allocation would accurately reflect nestling need. At eight nests we temporarily removed one nestling to deprive it of food, and video-taped the nestling and xii adult behaviors after the nestling was returned to the nest. Both male and female parents preferentially allocated food to the hungry nestling, but used different provisioning strategies and nestling cues to achieve this result. Males increased their visit rate and food load and used begging start order and initial beak height, as well as beg duration and intensity, as cues to allocate food. By contrast, females only increased food load and allocated resources to the nestling that begged more intensely. Most of our study nests were attended by only one parent. Hungry nestlings in male and female nests used different begging strategies associated with the cues employed by the parent (start/height in male nests, beg intensity in female nests). Nestlings appear to learn which begging signal components are most likely to generate food rewards. We suggest that conditioned
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