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Integrated principles of zoology PDF

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Fourteenth Hickman Edition About the cover: Something in the solitary majesty of the polar bear speaks to us despite the fact the Roberts predators often elicit less sense of kinship than do their prey. Polar bears hunt seals from Z sea ice. They feast on highly digestible seal fat which they store ahead of leaner times. Stored fat must last through the inter-ice period, at least four months, because the bears fast once the sea ice melts. The length of the fast is increasing. Global climate change Keen and ocean warming have resulted in earlier melt dates for sea ice: in the Canadian Arctic O melting occurs fi ve to eight days earlier each decade. By current estimates, sea ice now I breaks up three weeks earlier than in the 1970’s. Polar bears must swim longer distances n Larson to shore and so are more vulnerable to drowning. Increased stress and shorter feeding t seasons are particularly hard on pregnant females—they fast eight months because e Og birth occurs on shore when other bears return to r feeding. One polar bear population decreased 22% a I’Anson in the last twenty years, but human contact with t bears increased, presumably because bears have e d expanded their on-shore searches for food. What L the future holds for polar bears is hard to predict. P Eisenhour We can reduce our rates of bear harvest, but r whether we can slow the rate at which sea ice Oi n melts on a relevant time scale remains to be seen. c i Integrated Principles of Zoology p l Online Learning Center e G s Everything you need is online! Visit www.mhhe.com/hickmanipz14e for a wide array of o teaching and learning tools. Here is what you will fi nd: f • FOR INSTRUCTORS: Access to the new McGraw-Hill Presentation Center including Y all of the illustrations, photographs, and tables from Integrated Principles of Zoology; a password-protected Instructor’s Manual; PowerPoint lecture outlines; and other helpful resources for your course. • FOR STUDENTS: Online activities such as chapter quizzing, key term fl ash cards, Integ ra ted Princi ples of web links, and more! Hickman ZOOLOGY Roberts Keen Larson I’Anson Eisenhour Fourteenth Edition Cleveland P. Hickman, Jr. WASHINGTON AND LEE UNIVERSITY Larry S. Roberts FLORIDA INTERNATIONAL UNIVERSITY Susan L. Keen UNIVERSITY OF CALIFORNIA AT DAVIS Allan Larson WASHINGTON UNIVERSITY Helen I’Anson WASHINGTON AND LEE UNIVERSITY David J. Eisenhour MOREHEAD STATE UNIVERSITY Original Artwork by WILLIAM C. OBER, M.D. Washington and Lee University and Shoals Marine Laboratory and CLAIRE W. GARRISON, B.A. Shoals Marine Laboratory, Cornell University hhiicc7700004499__ffmm__ii--xxiivv..iinndddd ii 88//1100//0077 44::4422::5533 PPMM INTEGRATED PRINCIPLES OF ZOOLOGY, FOURTEENTH EDITION Published by McGraw-Hill, a business unit of The McGraw-Hill Companies, Inc., 1221 Avenue of the Americas, New York, NY 10020. Copyright © 2008 by The McGraw-Hill Companies, Inc. All rights reserved. Previous editions 2006, 2004, 2001, and 1997. No part of this publication may be reproduced or distributed in any form or by any means, or stored in a database or retrieval system, without the prior written consent of The McGraw-Hill Companies, Inc., including, but not limited to, in any network or other electronic storage or transmission, or broadcast for distance learning. Some ancillaries, including electronic and print components, may not be available to customers outside the United States. This book is printed on recycled, acid-free paper containing 10% postconsumer waste. 1 2 3 4 5 6 7 8 9 0 DOW/DOW 0 9 8 7 ISBN 978–0–07–297004–3 MHID 0–07–297004–9 Publisher: Janice Roerig-Blong Executive Editor: Patrick E. Reidy Developmental Editor: Debra A. Henricks Senior Marketing Manager: Tami Petsche Project Manager: April R. Southwood Senior Production Supervisor: Laura Fuller Lead Media Project Manager: Jodi K. Banowetz Media Producer: Daniel M. Wallace Associate Design Coordinator: Brenda A. Rolwes Cover Designer: Studio Montage, St. Louis, Missouri Senior Photo Research Coordinator: John C. Leland Photo Research: Mary Reeg Supplement Producer: Melissa M. Leick Compositor: Laserwords Private Limited Typeface: 10/12 Garamond Printer: R. R. Donnelley Willard, OH Front cover image: Polar Bear; © Digital Vision Back cover images: Polar bear paws; © Creatas/PunchStock, Polar bear standing on the ice; © Geostock/ Getty Images The credits section for this book begins on page 880 and is considered an extension of the copyright page. Library of Congress Cataloging-in-Publication Data Integrated principles of zoology / Cleveland P. Hickman, Jr. ... [et al.]. – 14th ed. p. cm. Includes index. ISBN 978–0–07–297004–3 — ISBN 0–07–297004–9 (hard copy : alk. paper) 1. Zoology. I. Hickman, Cleveland P. QL47.2.H54 2008 590–dc22 2007024506 www.mhhe.com hhiicc7700004499__ffmm__ii--xxiivv..iinndddd iiii 88//1100//0077 44::4433::1133 PPMM C O N T E N T S I N B R I E F About the Authors ix 20 Crustaceans 420 Preface xi 21 Hexapods 441 22 Chaetognaths, Echinoderms, and Hemichordates 469 23 Chordates 496 P A R T O N E 24 Fishes 514 25 Early Tetrapods and Modern Amphibians 543 Introduction to Living Animals 26 Amniote Origins and Nonavian Reptiles 563 1 Life: Biological Principles and the Science of Zoology 2 27 Birds 585 2 The Origin and Chemistry of Life 21 28 Mammals 612 3 Cells as Units of Life 37 4 Cellular Metabolism 58 P A R T F O U R P A R T T W O Activity of Life Continuity and Evolution of Animal Life 29 Support, Protection, and Movement 644 30 Homeostasis: Osmotic Regulation, Excretion, and Temperature 5 Genetics: A Review 76 Regulation 666 6 Organic Evolution 104 31 Internal Fluids and Respiration 686 7 The Reproductive Process 137 32 Digestion and Nutrition 708 8 Principles of Development 158 33 Nervous Coordination: Nervous System and Sense Organs 726 34 Chemical Coordination: Endocrine System 753 35 Immunity 771 P A R T T H R E E 36 Animal Behavior 785 Diversity of Animal Life 9 Architectural Pattern of an Animal 185 P A R T F I V E 10 Taxonomy and Phylogeny of Animals 199 11 Protozoan Groups 217 Animals and Their Environments 12 Sponges and Placozoans 246 37 The Biosphere and Animal Distribution 806 13 Radiate Animals 260 38 Animal Ecology 825 14 Flatworms, Mesozoans, and Ribbon Worms 289 15 Gnathiferans and Smaller Lophotrochozoans 313 16 Molluscs 331 Glossary 843 17 Annelids and Allied Taxa 362 Credits 880 Index 883 18 Smaller Ecdysozoans 384 19 Trilobites, Chelicerates, and Myriapods 402 iiiiii hhiicc7700004499__ffmm__ii--xxiivv..iinndddd iiiiii 88//1100//0077 44::4433::1133 PPMM T A B L E O F C O N T E N T S About the Authors ix CHAPTER 4 Preface xi Cellular Metabolism 58 Energy and the Laws of Thermodynamics 59 The Role of Enzymes 60 P A R T O N E Chemical Energy Transfer by ATP 62 Cellular Respiration 63 Metabolism of Lipids 70 Metabolism of Proteins 71 Management of Metabolism 72 Summary 73 P A R T T W O Introduction to Living Animals CHAPTER 1 Life: Biological Principles and the Science of Zoology 2 Fundamental Properties of Life 4 Zoology as a Part of Biology 10 Principles of Science 11 Theories of Evolution and Heredity 15 Continuity and Evolution of Animal Life Summary 19 CHAPTER 2 CHAPTER 5 The Origin and Chemistry of Life 21 Genetics: A Review 76 Water and Life 22 Mendel’s Investigations 77 Organic Molecular Structure of Living Systems 24 Chromosomal Basis of Inheritance 77 Chemical Evolution 28 Mendelian Laws of Inheritance 81 Origin of Living Systems 31 Gene Theory 90 Precambrian Life 32 Storage and Transfer of Genetic Information 91 Summary 35 Genetic Sources of Phenotypic Variation 100 Molecular Genetics of Cancer 101 CHAPTER 3 Summary 101 Cells as Units of Life 37 Cell Concept 38 Organization of Cells 40 Mitosis and Cell Division 52 Summary 56 iivv hhiicc7700004499__ffmm__ii--xxiivv..iinndddd iivv 88//1100//0077 44::4433::1144 PPMM www.mhhe.com/hickmanipz14e Table of Contents v CHAPTER 6 CHAPTER 9 Organic Evolution 104 Architectural Pattern of an Animal 185 Origins of Darwinian Evolutionary Theory 105 Hierarchical Organization of Animal Complexity 186 Darwinian Evolutionary Theory: The Evidence 108 Animal Body Plans 187 Revisions of Darwin’s Theory 126 Components of Metazoan Bodies 190 Microevolution: Genetic Variation and Change Within Complexity and Body Size 193 Species 126 Summary 195 Macroevolution: Major Evolutionary Events 132 Summary 134 CHAPTER 10 Taxonomy and Phylogeny of Animals 199 CHAPTER 7 The Reproductive Process 137 Linnaeus and Taxonomy 200 Species 201 Nature of the Reproductive Process 138 Taxonomic Characters and Phylogenetic Reconstruction 205 The Origin and Maturation of Germ Cells 142 Theories of Taxonomy 207 Reproductive Patterns 146 Major Divisions of Life 212 Structure of Reproductive Systems 147 Major Subdivisions of the Animal Kingdom 213 Endocrine Events That Orchestrate Reproduction 149 Summary 215 Summary 156 CHAPTER 11 CHAPTER 8 Protozoan Groups 217 Principles of Development 158 How Do We Defi ne Protozoan Groups? 218 Early Concepts: Preformation Versus Epigenesis 159 Form and Function 221 Fertilization 160 Major Protozoan Taxa 228 Cleavage and Early Development 162 Phylogeny and Adaptive Diversifi cation 243 An Overview of Development Following Cleavage 164 Summary 244 Suites of Developmental Characters 166 Mechanisms of Development 170 CHAPTER 12 Gene Expression During Development 172 Vertebrate Development 175 Sponges and Placozoans 246 Development of Systems and Organs 179 Summary 182 Origin of Metazoa 247 Phylum Porifera: Sponges 248 Phylum Placozoa 257 Summary 258 P A R T T H R E E CHAPTER 13 Radiate Animals 260 Phylum Cnidaria 261 Phylum Ctenophora 282 Phylogeny and Adaptive Diversifi cation 285 Summary 287 Diversity of Animal Life hhiicc7700004499__ffmm__ii--xxiivv..iinndddd vv 88//1100//0077 44::4433::2244 PPMM vi Table of Contents CHAPTER 14 Phylum Loricifera 395 Clade Panarthropoda 396 Flatworms, Mesozoans and Ribbon Worms 289 Phylogeny 399 Summary 400 Phylum Acoelomorpha 290 Clades Within Protostomia 291 CHAPTER 19 Phylum Platyhelminthes 292 Phylum Mesozoa 307 Trilobites, Chelicerates, and Myriapods 402 Phylum Nemertea (Rhynchocoela) 307 Summary 311 Phylum Arthropoda 403 Subphylum Trilobita 406 CHAPTER 15 Subphylum Chelicerata 407 Subphylum Myriapoda 414 Gnathiferans and Smaller Lophotrochozoans 313 Phylogeny and Adaptive Diversifi cation 416 Summary 418 Clade Gnathifera 314 Phylum Gnathostomulida 314 Phylum Micrognathozoa 316 CHAPTER 20 Phylum Rotifera 316 Crustaceans 420 Phylum Acanthocephala 319 Phylum Cycliophora 321 Subphylum Crustacea 422 Phylum Gastrotricha 321 A Brief Survey of Crustaceans 430 Phylum Entoprocta 323 Phylogeny and Adaptive Diversifi cation 439 Lophophorates 324 Summary 439 Phylum Ectoprocta (Bryozoa) 325 Phylum Brachiopoda 326 Phylum Phoronida 327 CHAPTER 21 Phylogeny 328 Hexapods 441 Summary 329 Class Insecta 443 CHAPTER 16 Insects and Human Welfare 459 Phylogeny and Adaptive Diversifi cation 465 Molluscs 331 Summary 466 Molluscs 332 Form and Function 333 CHAPTER 22 Classes of Molluscs 336 Phylogeny and Adaptive Diversifi cation 357 Chaetognaths, Echinoderms, and Hemichordates 469 Summary 360 Phylum Chaetognatha 471 Clade Ambulacraria 472 CHAPTER 17 Phylum Echinodermata 472 Annelids and Allied Taxa 362 Phylogeny and Adaptive Diversifi cation 488 Phylum Hemichordata 490 Phylum Annelida, Including Pogonophorans (Siboglinids) 364 Phylogeny and Adaptive Diversifi cation 493 Phylum Echiura 379 Summary 494 Phylum Sipuncula 380 Evolutionary Signifi cance of Metamerism 381 CHAPTER 23 Phylogeny and Adaptive Diversifi cation 381 Summary 382 Chordates 496 CHAPTER 18 The Chordates 497 Five Chordate Hallmarks 500 Smaller Ecdysozoans 384 Ancestry and Evolution 501 Subphylum Urochordata (Tunicata) 502 Phylum Nematoda: Roundworms 386 Subphylum Cephalochordata 504 Phylum Nematomorpha 393 Subphylum Vertebrata (Craniata) 505 Phylum Kinorhyncha 394 Summary 512 Phylum Priapulida 394 hhiicc7700004499__ffmm__ii--xxiivv..iinndddd vvii 88//1100//0077 44::4433::2299 PPMM www.mhhe.com/hickmanipz14e Table of Contents vii CHAPTER 24 P A R T F O U R Fishes 514 Ancestry and Relationships of Major Groups of Fishes 515 Living Jawless Fishes 515 Class Chondrichthyes: Cartilaginous Fishes 520 Osteichthyes: Bony Fishes 525 Structural and Functional Adaptations of Fishes 529 Summary 541 CHAPTER 25 Early Tetrapods and Modern Amphibians 543 Movement onto Land 544 Early Evolution of Terrestrial Vertebrates 544 Modern Amphibians 548 Summary 561 Activity of Life CHAPTER 26 CHAPTER 29 Amniote Origins and Nonavian Reptiles 563 Support, Protection, and Movement 644 Origin and Early Evolution of Amniotes 564 Characteristics of Nonavian Reptiles That Distinguish Them from Integument 645 Amphibians 568 Skeletal Systems 648 Characteristics and Natural History of Reptilian Orders 570 Animal Movement 654 Summary 583 Summary 663 CHAPTER 27 CHAPTER 30 Birds 585 Homeostasis: Osmotic Regulation, Excretion, and Temperature Regulation 666 Origin and Relationships 586 Structural and Functional Adaptations for Flight 587 Water and Osmotic Regulation 667 Flight 598 Invertebrate Excretory Structures 671 Migration and Navigation 601 Vertebrate Kidney 673 Social Behavior and Reproduction 603 Temperature Regulation 679 Bird Populations 606 Summary 684 Summary 610 CHAPTER 31 CHAPTER 28 Internal Fluids and Respiration 686 Mammals 612 Internal Fluid Environment 687 Origin and Evolution of Mammals 613 Composition of Blood 688 Structural and Functional Adaptations of Mammals 617 Circulation 690 Humans and Mammals 631 Respiration 698 Human Evolution 632 Summary 706 Summary 640 hhiicc7700004499__ffmm__ii--xxiivv..iinndddd vviiii 88//1100//0077 44::4433::2299 PPMM viii Table of Contents CHAPTER 32 P A R T F I V E Digestion and Nutrition 708 Feeding Mechanisms 709 Digestion 712 Organization and Regional Function of Alimentary Canals 714 Regulation of Food Intake 720 Nutritional Requirements 722 Summary 724 CHAPTER 33 Nervous Coordination: Nervous System and Sense Organs 726 Neurons: Functional Units of Nervous Systems 727 Synapses: Junctions Between Nerves 730 Evolution of Nervous Systems 733 Animals and Their Environments Sense Organs 740 Summary 751 CHAPTER 37 The Biosphere and Animal Distribution 806 CHAPTER 34 Chemical Coordination: Endocrine System 753 Distribution of Life on Earth 807 Animal Distribution (Zoogeography) 817 Mechanisms of Hormone Action 754 Summary 823 Invertebrate Hormones 756 Vertebrate Endocrine Glands and Hormones 758 CHAPTER 38 Summary 769 Animal Ecology 825 CHAPTER 35 The Hierarchy of Ecology 826 Immunity 771 Extinction and Biodiversity 839 Summary 841 Susceptibility and Resistance 772 Innate Defense Mechanisms 772 Immunity in Invertebrates 774 Glossary 843 Acquired Immune Response in Vertebrates 775 Credits 880 Blood Group Antigens 782 Index 883 Summary 783 CHAPTER 36 Animal Behavior 785 Describing Behavior: Principles of Classical Ethology 787 Control of Behavior 788 Social Behavior 792 Summary 802 hhiicc7700004499__ffmm__ii--xxiivv..iinndddd vviiiiii 88//1100//0077 44::4433::3377 PPMM A B O U T T H E A U T H O R S CLEVELAND P. HICKMAN, JR. Dr. Roberts has published many National Academies Education Fellow in research articles and reviews. He has the Life Sciences for 2005–2006. Her inter- Cleveland P. Hickman, Jr., Professor served as President of the American ests include weight training, horseback rid- Emeritus of Biology at Washington and Society of Parasitologists, Southwestern ing, gardening, travel, and mystery novels. Lee University in Lexington, Virginia, has Association of Parasitologists, and South- Dr. Keen can be contacted at: taught zoology and animal physiology for eastern Society of Parasitologists, and is a [email protected] more than 30 years. He received his Ph.D. member of numerous other professional in comparative physiology from the Uni- societies. Dr. Roberts also serves on the versity of British Columbia, Vancouver, ALLAN LARSON Editorial Board of the journal, P arasitology B.C., in 1958 and taught animal physiol- Research. His hobbies include scuba div- Allan Larson is a professor at Washington ogy at the University of Alberta before ing, underwater photography, and tropical University, St. Louis, MO. He received his moving to Washington and Lee Univer- horticulture. Ph.D. in genetics at the University of Cali- sity in 1967. He has published numer- Dr. Roberts can be contacted at: fornia, Berkeley. His fi elds of specializa- ous articles and research papers in fi sh [email protected] tion include evolutionary biology, molec- physiology, in addition to co-authoring ular population genetics and systematics, these highly successful texts: Integrated and amphibian systematics. He teaches Principles of Zoology, Biology of Animals, SUSAN KEEN courses in introductory genetics, zoology, Animal Diversity, Laboratory Studies in Susan Keen is a lecturer in the Section of macroevolution, molecular evolution, and Animal Diversity, and Laboratory Studies Evolution and Ecology at the University of the history of evolutionary theory, and in Integrated Principles of Zoology. California at Davis. She received her Ph.D. has organized and taught a special course Over the years Dr. Hickman has led in zoology from the University of Califor- in evolutionary biology for high-school many fi eld trips to the Galápagos Islands. nia at Davis, following a M.Sc. from the teachers. His current research is on intertidal zona- University of Michigan at Ann Arbor. She Dr. Larson has an active research labo- tion and marine invertebrate systematics is a native of Canada and obtained her ratory that uses DNA sequences to exam- in the Galápagos. He has published three undergraduate education at the University ine evolutionary relationships among ver- field guides in the Galápagos Marine of British Columbia in Vancouver. tebrate species, especially in salamanders Life Series for the identifi cation of echi- Dr. Keen is an invertebrate zoologist and lizards. The students in Dr. Larson’s noderms, marine molluscs, and marine fascinated with jellyfi sh life histories. She laboratory have participated in zoological crustaceans. has a particular interest in life cycles where fi eld studies around the world, including His interests include scuba diving, both asexual and sexual phases of organ- projects in Africa, Asia, Australia, Mada- woodworking, and participating in cham- isms are present, as they are in most jel- gascar, North America, South America, ber music ensembles. lyfi shes. Her other research has included the Indo-Pacific Ocean, and the Carib- Dr. Hickman can be contacted at: work on sessile marine invertebrate com- bean Islands. Dr. Larson has authored [email protected] munities, spider populations, and Andean numerous scientific publications, and potato evolution. has edited for the journals The American Dr. Keen has been teaching evolution Naturalist, Evolution, Journal of Experi- LARRY S. ROBERTS and animal diversity within the Introduc- mental Zoology, Molecular Phylogenetics Larry S. Roberts, Professor Emeritus of tory Biology series for 13 years. She enjoys and Evolution, and Systematic Biology. Biology at Texas Tech University and an all facets of the teaching process, from Dr. Larson serves as an academic advisor adjunct professor at Florida International lectures and discussions to the design of to undergraduate students and supervises University, has extensive experience effective laboratory exercises. In addition the undergraduate biology curriculum at teaching invertebrate zoology, marine to her work with introductory biology, Washington University. biology, parasitology, and developmental she offers seminars for the Davis Honors Dr. Larson can be contacted at: biology. He received his Sc.D. in parasitol- Challenge program, and for undergraduate [email protected] ogy at the Johns Hopkins University and is and graduate students interested in teach- the lead author of Schmidt and Roberts’s ing methods for biology. She was given HELEN I’ANSON Foundations of Parasitology, sixth edition. an Excellence in Education Award from Dr. Roberts is also co-author of I ntegrated the Associated Students group at Davis Helen I’Anson, a native of England, is Principles of Zoology, Biology of Animals, in 2004. She attended the National Acad- professor of biology at Washington and and A nimal Diversity, and is author of The emies Summer Institute on Undergraduate Lee University in Lexington, Virginia. She Underwater World of Sport Diving. Education in Biology in 2005, and was a received her Ph.D. in physiology at the ix hhiicc7700004499__ffmm__ii--xxiivv..iinndddd iixx 88//1100//0077 44::4433::4433 PPMM

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