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Committee on Nutrient Requirements of Small Ruminants Board on Agriculture and Natural Resources Division on Earth and Life Studies

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THE NATIONAL ACADEMIES PRESS 500 Fifth Street, N.W. Washington, DC 20001 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. This study was supported by grants from the National Sheep Industry Improvement Center (U.S. Department of Agriculture), the Joe Skeen Institute for Rangeland Restoration (Montana State University, New Mexico State University, and Texas A&M University), the American Sheep Industry Association, the Montana Sheep Institute, internal National Research Council funds, and general support of the Animal Nutrition series provided by the Department of Health and Human Services (U.S. Food and Drug Administration) under Award No. 223-01-01-2460. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the views of the organizations or agencies that provided support for the project. Library of Congress Cataloging-in-Publication Data Nutrient requirements of small ruminants : sheep, goats, cervids, and New World camelids / Committee on Nutrient Requirements of Small Ruminants, Board on Agriculture and Natural Resources, Division on Earth and Life Studies, The National Research Council. p. cm. Includes bibliographical references and index. ISBN 0-309-10213-8 (cloth) -- 978-0-309-10213-1 (cloth) 1. Ruminants--Feeding and feeds. 2. Ruminants-- Nutrition--Requirements. I. National Research Council (U.S.). Committee on Nutrient Requirements of Smal Ruminants. SF95.N78 2006 636.30852--dc22 2006028306 Additional copies of this report are available from the National Academies Press, 500 Fifth Street, N.W., Lockbox 285, Washington, DC 20055; (800) 624-6242 or (202) 334-3313 (in the Washington metropolitan area); Internet, http://www.nap.edu Copyright 2007 by the National Academy of Sciences. All rights reserved. Printed in the United States of America

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The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distin- guished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the authority of the char- ter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Ralph J. Cicerone is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is auton- omous in its administration and in the selection of its members, sharing with the National Acad- emy of Sciences the responsibility for advising the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. Wm. A. Wulf is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to se- cure the services of eminent members of appropriate professions in the examination of policy matters pertaining to the health of the public. The Institute acts under the responsibility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and educa- tion. Dr. Harvey V. Fineberg is president of the Institute of Medicine. The National Research Council was organized by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy's purposes of furthering knowledge and advising the federal government. Functioning in accordance with gen- eral policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in provid- ing services to the government, the public, and the scientific and engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Ralph J. Cicerone and Dr. Wm. A. Wulf are chair and vice chair, respectively, of the National Research Council. www.national-academies.org

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COMMITTEE ON NUTRIENT REQUIREMENTS OF SMALL RUMINANTS JAMES E. (ED) HUSTON, Chair, Texas A&M University, San Angelo ROBERT G. WHITE, Vice-Chair, University of Alaska, Fairbanks BRIAN J. BEQUETTE, University of Maryland, College Park HUGH DOVE, Commonwealth Scientific and Industrial Research Organisation, Canberra, Australia ARTHUR L. GOETSCH, Langston University, Langston, Oklahoma BRET W. HESS, University of Wyoming, Laramie MAXIMINO HUERTA BRAVO, Universidad Autnoma Chapingo, Texcoco, Mexico DAVID G. PUGH, Fort Dodge Animal Health, Waverly, Alabama SANDRA G. SOLAIMAN, Tuskegee University, Alabama Staff AUSTIN J. LEWIS, Study Director JAMIE S. JONKER, Study Director* DONNA LEE JAMEISON, Senior Project Assistant** RUTH S. ARIETI, Project Assistant PEGGY TSAI, Research Associate External Support PAULA T. WHITACRE, Editor *Through June 2004 **Through January 2006 v

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BOARD ON AGRICULTURE AND NATURAL RESOURCES W. REG GOMES, Chair, University of California, Oakland SANDRA J. BARTHOLMEY, University of Illinois at Chicago ROGER N. BEACHY, Donald Danforth Plant Science Center, St. Louis, Missouri H. H. CHENG, University of Minnesota, St. Paul BRUCE L. GARDNER, University of Maryland, College Park, Maryland JEAN HALLORAN, Consumer Policy Institute/Consumers Union, Yonkers, New York HANS R. HERREN, Millennium Institute, Arlington, Virginia KIRK C. KLASING, University of California, Davis BRIAN W. MCBRIDE, University of Guelph, Guelph, Canada TERRY L. MEDLEY, E. I. duPont de Nemours & Co., Wilmington, Delaware ROBERT PAARLBERG, Wellesley College, Watertown, Massachusetts ALICE N. PELL, Cornell University, Ithaca, New York BOBBY PHILLS, Florida A&M University, Tallahassee SONYA B. SALAMON, University of Illinois, Urbana-Champaign HAL SALWASSER, Oregon State University, Corvallis, Oregon PEDRO A. SANCHEZ, The Earth Institute at Columbia University, Palisades, New York B. L. TURNER, II, Clark University, Worcester, Massachusetts LAURIAN UNNEVEHR, University of Illinois at Urbana-Champaign TILAHUN D. YILMA, University of California, Davis JAW-KAI WANG, University of Hawaii, Honolulu Staff ROBIN A. SCHOEN, Director KAREN L. IMHOF, Administrative Assistant AUSTIN J. LEWIS, Program Officer PEGGY TSAI, Research Associate RUTH S. ARIETI, Project Assistant vi

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Preface Small ruminants, especially sheep and goats, are an im- elk [Cervus elaphus], and caribou/reindeer [Rangifer taran- portant part of the economic and social life of many nations dus]), and New World camelids (alpacas [Lama glama] and throughout the world. Numerous other species of small ru- llamas [Vicugna pacos]). The second important decision was minants range the Earth in undomesticated herds, some of how to organize the publication--by species or by topic, which are managed and others which are not. Information such as individual nutrients. Ultimately, the committee de- about the dietary habits and nutrient requirements of these cided that to organize the report by species would result in animals is crucial in managing their wellbeing and in con- a large amount of repetition and redundancy. Therefore, the tributing to the livelihoods of people who depend on them. report is organized by topic. Thus, for example, there are Proper nutrition and management of small ruminants is also chapters on individual nutrients, and the species are covered essential for the maintenance of various natural ecosystems within each chapter. The committee recognized that this from the Arctic Circle to the Sahara Desert. makes the publication somewhat less convenient for readers The National Research Council has published reports on interested in one species only. To address this, tables of nu- the Nutrient Requirements of Sheep (6th edition, 1985) and trient requirements are organized by species, and one chap- the Nutrient Requirements of Goats (1st edition, 1981), but ter discusses practical feeding and nutrition on a species-by- a large amount of new information has accumulated during species basis. the last 25 years. Other small ruminants have not been ad- This report represents the concerted efforts of all com- dressed in the Nutrient Requirements series. This new pub- mittee members. The committee was appointed in April lication updates the sheep and goat publications and adds in- 2004 and held its first meeting in May 2004. After two more formation about other small ruminant species. In addition, in-person meetings and several teleconferences, an initial information about New World camelids (alpacas and llamas) draft was sent to reviewers in November 2005. The commit- is included to the extent that it is available. tee hopes the publication will serve as an important refer- The committee appointed to write this report faced two ence for all concerned with the nutrition, feeding, health, important decisions at the outset. The first decision was and welfare of small ruminants and that it will also serve to which species to include as "small ruminants." After consid- stimulate more research on this important group of animals. ering various factors such as body size, amount of scientific research data, absence of alternative sources of information, and general interest of potential users, the committee de- ED HUSTON cided to include the following species: sheep (Ovis aries), Chair, Committee on goats (Capra hircus), cervids (white-tailed deer [Odocoileus Nutrient Requirements of virginianus], red deer [Cervus elaphus], wapiti/American Small Ruminants vii

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Acknowledgments This report has been reviewed in draft form by persons Although the reviewers listed above have provided con- chosen for their diverse perspectives and technical expertise structive comments and suggestions, they were not asked to in accordance with procedures approved by the National endorse the conclusions or recommendations, nor did they Research Council's Report Review Committee. The purpose see the final draft of the report before its release. The review of this independent review is to provide candid and critical of this report was overseen by Michael L. Galyean, Texas comments that will assist the institution in making its pub- Tech University. Appointed by the National Research Coun- lished report as sound as possible and to ensure that the re- cil, he was responsible for making certain that an independ- port meets institutional standards of objectivity, evidence, ent examination of this report was carried out in accordance and responsiveness to the study charge. The review com- with institutional procedures and that all review comments ments and draft manuscript remain confidential to protect were carefully considered. Responsibility for the final con- the integrity of the deliberative process. We wish to thank tent of this report rests entirely with the author committee the following individuals for their review of this report: and the institution. We are grateful for contributions of Larry Miller, CSREES- Antonello Cannas, University of Sassari, Sardinia, Italy USDA, who has been a good friend to the industries that will George F. W. Haenlein, University of Delaware, Newark, be served by this report. Larry is an ever-present entity when- DE (retired) ever the animal industries, the scientific community, and U.S. Patrick G. Hatfield, Montana State University, Bozeman, MT governmental agencies interact and always a facilitator, a role Robert J. Hudson, University of Alberta, Edmonton, that he fulfilled in the production of this document. Alberta, Canada Members of the author committee express appreciation to Lyndon N. Irwin, Southwest Missouri State University, the National Research Council and parent agency, The Springfield, MO National Academies, for entrusting us to prepare this greatly Woody Lane, Lane Livestock Services, Roseburg, OR anticipated and respected document. Special thanks are given Jean-Marie Luginbuhl, North Carolina State University, to Charlotte Kirk Baer, former Director of the Board on Raleigh, NC Agriculture and Natural Resources, for her initiation of this Daniel G. Morrical, Iowa State University, Ames, IA project and early guidance, and to Robin Schoen, current John V. Nolan, University of New England, Armidale, Director, for her administrative oversight of this process and NSW, Australia continued encouragement. A deeply felt thank you goes to Fredric N. Owens, Pioneer Hi-Bred International, Donna Lee Jameison, Senior Project Assistant, and to Ruthie West Des Moines, IA Arieti, Project Assistant, for their technical contributions in Rodney L. Preston, Texas Tech University, Lubbock, TX monitoring, formatting, editing, posting, and other responsi- (retired) bilities, and for their extremely good nature while perform- Duane E. Ullrey, Michigan State University, East Lansing, ing these vital services. Finally, the committee wishes to MI (retired) thank Austin Lewis, Study Director, who led us on a daily Larry W. Varner, Purina Mills Inc, Seguin, TX basis by giving us information, feedback, critique, and encouragement and brought us to successful completion. ix

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Contents SUMMARY 1 1 INTRODUCTION 3 2 ANATOMY, DIGESTIVE PHYSIOLOGY, AND NUTRIENT UTILIZATION 5 3 INGESTIVE BEHAVIOR AND FEED INTAKE 26 4 ENERGY 39 5 PROTEIN 81 6 ESSENTIAL FATTY ACIDS 106 7 MINERALS 112 8 VITAMINS 150 9 WATER 173 10 PLANT FACTORS AFFECTING NUTRIENT AVAILABILITY 189 11 NUTRIENT SOURCES AND FEEDING PRACTICES 201 12 ENVIRONMENTAL IMPACT 216 13 NUTRITIONAL AND METABOLIC DISEASES, AILMENTS, AND MODIFIERS 220 14 PROVIDING REQUIRED NUTRIENTS TO SMALL RUMINANTS 232 NUTRIENT REQUIREMENT TABLES 244 Nutrient Requirements of Sheep, 244 Nutrient Requirements of Goats, 271 Nutrient Requirements of Cervids, 300 Nutrient Requirements of New World Camelids, 308 xi

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xii CONTENTS FEED COMPOSITION TABLES 312 Common Feedstuffs, 312 Novel Feedstuffs, 322 Mineral Supplements, 332 OTHER TABLES 335 Conversion Factors, SI Prefixes, and Termperature Conversions, 335 APPENDIXES A COMMITTEE STATEMENT OF TASK 337 B ABBREVIATONS AND ACRONYMS 338 C COMMITTEE MEMBER BIOGRAPHIES 342 D BOARD ON AGRICULTURE AND NATURAL RESOURCES PUBLICATIONS 344 INDEX 347

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Tables and Figures TABLES 2-1 Digestible Nitrogen Intake and Urea Synthesis and Recycling in DifferentAnimal Species (g N/day), 19 3-1 Selected Comparisons of Diet Compositions of Small Ruminants, 28 3-2 Potential Intakes (kg DM/day) of Twin-Suckling Ewes (Mature Weight 60 kg; Relative Size 0.85), when Eating Grass Diets of 8, 10, or 12 MJ ME/kg DM during Lactation, 34 4-1 Predicted Empty Body Concentrations of Fat, Protein, and Energy in Sheep from Body Condition Score, 44 4-2 Energy and Protein Requirements of Small Ruminants for Pregnancy, 53 4-3 References, Number of Treatment Mean Observations, and Genotypes in the Database Used to Evaluate Prediction of Average Daily Gain, 55 4-4 Description of the Database Used to Evaluate Prediction of Average Daily Gain, 55 4-5 References, Number of Treatment Mean Observations, and Genotypes in the Database Used to Evaluate Prediction of Net Energy for Lactation and Average Daily Gain, 61 4-6 Description of the Database Used to Evaluate Prediction of Average Daily Gain and Net Energy for Lactation, 61 4-7 Adjustment Factors, 67 5-1 EssentialAminoAcid Composition (grams of amino acid per 100 grams protein) of Mixed Microbial Protein Compared with Carcass Tissue, Milk Casein, Skin, and Wool, 88 5-2 Wool Protein Composition (grams of amino acid per 100 grams protein), 89 5-3 References, Number of Treatment Mean Observations, and Genotypes in the Database Used to Evaluate Metabolizable Protein Requirements of the CNCPS-S Method, 98 5-4 Description of the Database Used to Evaluate Metabolizable Protein (MP) Requirements of the CNCPS-S Method, 99 7-1 Copper Absorption Based on Normal Molybdenum and Sulfur Content of the Diet, 128 9-1 Comparison of Estimates of Daily Water Requirements of Goats, 184 10-1 Summary of Some Observed Positive and Negative Effects of Plant Secondary Metabo- lites after Consumption by Mammalian Herbivores, 191 10-2 Fractionation of Forage Organic Matter According to the Detergent System, 194 10-3 Estimated Digestibility of Neutral Detergent Solubles in a Range of Grass Species, 196 11-1 Long-Chain Fatty Acids in Some Feedstuffs Fed to Small Ruminants, 204 15-1 Nutrient Requirements of Sheep (mature ewes and rams and yearlings maintenance and lactation), 246 xiii

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xiv TABLES AND FIGURES 15-2 Nutrient Requirements of Sheep (growing, finishing lambs, and yearlings during growth/gestation), 256 15-3 Mineral Requirements of Sheep, 266 15-4 Nutrient Requirements of Meat and Milk Goats (maintenance-gestation-lactation), 272 15-5 Nutrient Requirements of Goats (growing kids), 281 15-6 Nutrient Requirements of Goats (Angora), 288 15-7 Mineral Requirements of Meat and Milk Goats, 293 15-8 Mineral Requirements of Goats (Angora), 297 15-9 Nutrient Requirements of Cervid Species (white-tailed deer, reindeer/caribou, red deer and elk/wapiti), 301 15-10 Nutrient Requirements of New World Camelids, 309 15-11 Composition of Common Feedstuffs, 312 15-12 Composition of Pasture or Range Forage, Browse, and Other Novel Feedstuffs, 322 15-13 Compositions of Inorganic Mineral Sources on a 100% Dry Matter Basis, 332 15-14 Table of Conversion Factors, 335 FIGURES 2-1 Drawings of the gastrointestinal tracts of the sheep (a grass/roughage eater morphologi- cal feeding type) and the llama, 6 2-2 A drawing of the red deer (an intermediate morphological feeding type) showing the stomach in situ and isolated, 7 2-3 A drawing of the right aspect of the stomach of the goat (an intermediate morphological feeding type), 7 2-4 Photograph of the four stomach compartments of a yearling white-tailed deer (a concen- trate selector morphological feeding type), 8 2-5 The inside structure of the four stomach compartments of a white-tailed deer (a concen- trate selector morphological feeding type), 8 2-6 Drawings of the three stomach compartments of the llama, 9 3-1 Influence of standard reference weight of sheep (mature size when in average condition score) and relative size (current weight as a proportion of mature weight) on predicted intakes of an all-grass diet of nonlimiting digestibility (> 0.8), 33 3-2 Influence of relative size (proportion of mature live weight) on predicted DM intakes, ex- pressed in relation to mature size, 33 3-3 Influence of time since lambing on the predicted DM intake (% of standard reference weight) of ewes of relative size 0.85 consuming an all-grass diet and suckling either sin- gle or twin lambs, 34 3-4 Pasture intakes (kg DM/day) predicted for sheep of standard reference weight 50 kg, grazing improved grass pastures differing in average digestibility, 35 4-1 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the NRC method for growing/finishing sheep, 57 4-2 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the CNCPS-S method for growing/finishing sheep, 57 4-3 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the NRC method for growing finishing/sheep, 58 4-4 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the CNCPS-S method for growing/finishing sheep, 59 4-5 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the CNCPS-S method for lactating sheep, 62 4-6 The relationship between observed ADG (ADGobs) and ADG predicted (ADGpred) by the CNCPS-S method for lactating sheep, 63 4-7 The relationship between observed NEl (NEl-obs) and NEl predicted (NEl-pred) by the CNCPS-S method for lactating sheep, 63

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TABLES AND FIGURES xv 4-8 The relationship between observed NEl (NEl-obs) and NEl predicted (NEl-pred) by the CNCPS-S method for lactating sheep, 64 5-1 The relationship between metabolizable protein (MP) available (based on the difference between predicted metabolizable protein intake and metabolizable protein used for main- tenance; MPg-avail) and used for gain or maintenance (based on predicted metabolizable protein accreted in tissue or mobilized and used for maintenance; MPg-used), 91 5-2 The relationship between rumen degraded intake protein (DIP) as a percent of total di- gestible nutrients (TDN) and average daily gain (ADG) relative to body weight (BW)0.75 with a database of treatment mean observations from the literature for growing goats, 95 5-3 The relationship between rumen degraded intake protein (DIP) as a percent of total di- gestible nutrients (TDN) and average daily gain (ADG) relative to body weight (BW)0.75 with a database of treatment mean observations from the literature for mature goats, 96 5-4 The relationship between rumen degraded intake protein (DIP) as a percent of total di- gestible nutrients (TDN) and dry matter intake (DMI) relative to body weight (BW)0.75 with a database of treatment mean observations from the literature for growing goats, 96 5-5 The relationship between rumen degraded intake protein (DIP) as a percent of total di- gestible nutrients (TDN) and dry matter intake (DMI) relative to body weight (BW)0.75 with a database of treatment mean observations from the literature for mature goats, 96 7-1 Selenium requirements of growing lambs based on performance and selenium content of carcass, 136 10-1 Relationship between plant carbohydrate fractions and analytical components used to de- scribe them, 194 10-2 Typical changes in dry matter digestibility of a range of pasture grasses (upper panel) and legumes (lower panel) as spring progresses, 195 10-3 Changes in the digestibility of different plant parts of annual grass pasture during spring- autumn, 195 11-1 Annual weight change for single-bearing ewe, 208

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