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The Maned Wolves of Noel Kempff Mercado National Park

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The Maned Wolves of Noel Kempff Mercado National Park Smithsonian Institution Scholarly Press smithsonian contributions to zoology • number 639 Smithsonian Institution Scholarly Press TheA Chronology Maned Wolves of of MiddleNoel Kempff Missouri Mercado Plains National Park Village Sites Edited by Louise H. Emmons By Craig M. Johnson with contributions by Stanley A. Ahler, Herbert Haas, and Georges Bonani SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of “diffusing knowledge” was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: “It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge.” This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Botany Smithsonian Contributions to History and Technology Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Museum Conservation Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology In these series, the Institution publishes small papers and full-scale monographs that report on the research and collections of its various museums and bureaus. The Smithsonian Contributions Series are distributed via mailing lists to libraries, universities, and similar institu- tions throughout the world. Manuscripts submitted for series publication are received by the Smithsonian Institution Scholarly Press from authors with direct affilia- tion with the various Smithsonian museums or bureaus and are subject to peer review and review for compliance with manuscript preparation guidelines. General requirements for manuscript preparation are on the inside back cover of printed volumes. For detailed submissions require- ments and to review the “Manuscript Preparation and Style Guide for Authors,” visit the Submissions page at www.scholarlypress.si.edu. smithsonian contributions to zoology • number 639 The Maned Wolves of Noel Kempff Mercado National Park Edited by Louise H. Emmons WASHINGTON D.C. 2012 ABSTRACT Emmons, Louise H., editor. The Maned Wolves of Noel Kempff Mercado National Park. Smithsonian Contributions to Zool- ogy, number 639, xii + 135 pages, 60 figures, 37 tables, 2012.—We studied the behavioral ecology of maned wolves (Canidae: Chrysocyon brachyurus) for 10 years in Cerrado habitat of Noel Kempff National Park, Bolivia. Most data were collected by GPS-collar technology, which yielded over 37,000 locations in 27 collar deployments on 10 individuals. The eight chapters introduce the study area and methods (1) and describe daily and seasonal activity (2); movements and ranges (3); diet and en- ergetics (4); social interactions and reproduction (5); disease exposure, morbidity and mortality (6); maned wolf conservation (7); and finally, we synthesize the results in an overview of maned wolf behavioral ecology, with hypotheses about the unique form and function of this atypical canid (8). Activity was temperature related and sharply nocturnal in the dry season but partly diurnal in the rainy season. Adult home ranges were 40–123 km2, with strong seasonal variations in land use. Maned wolves averaged 14 km/night travel in dry months and 7 km/night during wet months. Breeding pairs shared territories with contigu- ous borders, which did not overlap with neighboring pairs. Young females twice stayed until adulthood on natal territories, as presumed helpers, and acquired the territory upon disappearance/death of the adult females. Females were the holders of territories into which males moved to form pairs. Young males all emigrated. By 8 years old, maned wolves showed extreme tooth wear, and dental disease was a major cause of morbidity. Habitat loss is the chief conservation issue for the species, but drought-related resource loss appears to be reducing the study area population. RESUMEN Emmons, Louise H., editor. Los Lobos de Crin de Parque Nacional Noel Kempff Mercado. Smithsonian Contributions to Zoology, número 639, xii + 135 páginas, 60 figuras, 37 tablas, 2012.—Estudiamos la ecología de los lobos de crin (Canidae: Chrysocyon) durante 10 años en el Parque Nacional Noel Kempff, Bolivia. La mayoría de los datos fueron colectados por la tecnología GPS-de- collar, que generó más de 37.000 ubicaciones en 10 individuos. Los ocho capítulos dan a conocer el área de estudio y métodos (1), y describen la actividad diaria y estacional (2), movimientos y rangos (3), dieta y energética (4), interacciones sociales y reproducción (5), exposición a enfermedades, morbilidad y mortalidad (6), conservación del lobo de crin (7) y, finalmente, sintetizamos los resultados con información general de la ecología y comportamiento del lobo de crin, con hipótesis acerca de la singular forma y función de este atípico cánido. La actividad estuvo relacionada con la temperatura y fue claramente nocturna en la estación seca, aunque en parte diurna en la temporada de lluvias. El área del territorio de adultos fue de 40–123 km2, con marcadas variaciones estacionales en el uso del hábitat. El desplazamiento promedio fue de 14 km/noche en los meses secos y 7 km durante los meses húmedos. Las parejas reproductoras compartieron territorios con límites contiguos, los cuales no se superpusieron con el de las parejas vecinas. Una hembra joven permaneció en dos ocasiones hasta la edad adulta en su territorio natal, se supone como “helper” y adquirió el territorio, a la desaparición/muerte de las hembras adultas. Las hembras fueron las dueñas de los territorios, en los que los machos se trasladaron para formar parejas. Los machos jóvenes emigraron. A los 8 años de edad, los lobos de crin mostraron desgastes extremos de los dientes, y las enfermedades dentales fueron una importante causa de morbilidad. La pérdida de hábitat es la amenaza principal para la conser- vación de la especie, pero la sequía relacionada con la pérdida de recursos parece estar reduciendo las poblaciones del área de estudio. Cover images from L to R: Maned wolf male M3 sitting in long grass; view during the wet season of the great Huanchaca tableland, a major feature of Noel Kempff Mercado National Park; and maned wolf youngster F9 with mane ruffled. All photos: L. H. Emmons. Published by Smithsonian Institution Scholarly Press P.O. Box 37012, MRC 957 Washington, D.C. 20013-7012 www.scholarlypress.si.edu Compilation copyright © 2012 Smithsonian Institution “Introduction: Study Site and Methods” and “Synthesis,” both by Louise H. Emmons, as well as the Preface and Acknowledg- ments are in the public domain. The rights to all other text and images in this publication, including cover and interior designs, are owned either by the Smithsonian Institution, by contributing authors, or by third parties. Fair use of materials is permitted for personal, educational, or noncommercial purposes. Users must cite author and source of content, must not alter or modify copyrighted content, and must comply with all other terms or restrictions that may be applicable. Users are responsible for securing permission from a rights holder for any other use. Library of Congress Cataloging-in-Publication Data The maned wolves of Noel Kempff Mercado National Park / edited by Louise H. Emmons. p. cm. -- (Smithsonian contributions to zoology ; NUMBER 639) Includes bibliographical references. 1. Maned wolf—Bolivia—Parque Nacional Noel Kempff Mercado. I. Emmons, Louise. QL737.C22.M3525 2012 599.77—dc23 2011045004 ISSN: 0081-0282 (print); 1943-6696 (online) The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48–1992. Contents LIST OF FIGURES v LIST OF TABLES vii PREFACE ix ACKNOWLEDGMENTS xi 1 INTRODUCTION: STUDY SITE AND METHODS 1 Louise H. Emmons 2 ACTIVITY Patterns 15 Louise H. Emmons, Verónica Chávez, Luis Fernando Del Aguila, and Sixto Angulo 3 RANGING Patterns 25 Louise H. Emmons, Verónica Chávez, Luis Fernando Del Aguila, Sixto Angulo, and Matthew J. Muir 4 Variation IN DIET AND RESOURCES 37 José Miguel Castro and Louise H. Emmons 5 SOCIAL INTERACTIONS AND REPRODUCTION 57 Louise H. Emmons, Verónica Chávez, Luis Fernando Del Aguila, Sixto Angulo, and Matthew J. Muir 6 MORBIDITY AND Mortality 77 Sharon L. Deem, Ellen Bronson, Sixto Angulo, Veronica Acosta, Suzan Murray, Richard G. Robbins, Urs Giger, Bruce Rothschild, and Louise H. Emmons 7 CONSERVation 91 Matthew J. Muir and Louise H. Emmons 8 SYNTHESIS 117 Louise H. Emmons REFERENCES 125 INDEX 133 Figures Frontispiece. Maned wolf male M8 on release with a new collar xii 1.1 Simplified diagram of major biomes of South America 2 1.2 Location of the study area in Bolivia 6 1.3 Campo Cerrado, Los Fierros savanna 7 1.4 Termite savanna (pampa) of Los Fierros savanna from the air 8 1.5 Maned wolf F3 in termite pampa 9 1.6 False color image of flooding levels, Los Fierros savanna 10 1.7 Bajío savanna at the beginning of February flooding 10 1.8 Mean monthly rainfall and total annual rainfall 11 1.9 Rainfall trends by season during the study years 11 1.10 Fires of 2009 12 2.1 Average proportion of “active” intervals between hourly fixes 17 2.2 Activity and collar temperature for F3 17 2.3 Hourly
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