611 150 138MB
English Pages 1856 [930] Year 2021
INTEGR ATED PRINC IPLES OF
EIGHTEENTH EDITION Cleveland P. Hickman, Jr. PR OFE SSOR E MER I T US WASHINGT ON AND LEE UNI VER SI T Y
Susan L. Keen UNI VER SI T Y OF C ALIFORNI A–DAV IS
David J. Eisenhour MOREHE AD STAT E UNI VER SI T Y
Allan Larson WASHINGT ON UNI VER SI T Y
Helen I’Anson WASHINGT ON AND LEE UNI VER SI T Y
Original Artwork by WILLIAM C. OBER, M.D. WASHINGTON AND LEE UNIVERSITY and CLAIRE W. OBER, B.A. WASHINGTON AND LEE UNIVERSITY
INTEGRATED PRINCIPLES OF ZOOLOGY Published by McGraw-Hill Education, 2 Penn Plaza, New York, NY 10121. Copyright © 2020 by McGraw-Hill Education. All rights reserved. Printed in the United States of America. Previous editions © 2017, 2014, and 2011. 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 McGraw-Hill Education, 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 acid-free paper. 1 2 3 4 5 6 7 8 9 LWI 21 20 19 ISBN 978-1-260-56597-3 MHID 1-260-56597-1 Cover Image: ©Shutterstock/idreamphoto
All credits appearing on page are considered to be an extension of the copyright page. The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not indicate an endorsement by the authors or McGraw-Hill Education, and McGraw-Hill Education does not guarantee the accuracy of the information presented at these sites.
mheducation.com/highered
CONTENTS IN BRIEF
20 Crustaceans
About the Authors ix Preface x
21 Hexapods
428
450
22 Chaetognaths, Echinoderms, and Hemichordates 480 23 Chordates
PART ONE
24 Fishes
Introduction to Living Animals 1 Life: Biological Principles and the Science of Zoology 1 2 The Origin and Chemistry of Life 20 3 Cells as Units of Life 36
506
524
25 Early Tetrapods and Modern Amphibians 552 26 Amniote Origins and Nonavian Reptiles 571 27 Birds
593
28 Mammals
619
4 Cellular Metabolism 57
PART FOUR PART T WO
Activity of Life
Continuity and Evolution of Animal Life
29 Support, Protection, and Movement 648
5 Genetics: A Review 73
30 Homeostasis: Osmotic Regulation, Excretion, and Temperature Regulation 669
6 Organic Evolution 103
31 Homeostasis: Internal Fluids and Respiration 689
7 The Reproductive Process 137
32 Digestion and Nutrition 711
8 Principles of Development 158
33 Nervous Coordination: Nervous System and Sense Organs 729 34 Chemical Coordination: Endocrine System 755
PART THREE
35 Immunity
773
36 Animal Behavior
787
Diversity of Animal Life 9 Architectural Pattern of an Animal 187 10 Taxonomy and Phylogeny of Animals 201 11 Unicellular Eukaryotes
220
PART FIVE
12 Sponges and Placozoans 250
Animals and Their Environments
13 Cnidarians and Ctenophores 264
37 Animal Distributions
14 Xenacoelomorpha, Platyzoa, and Mesozoa 294
38 Animal Ecology
806
826
15 Polyzoa and Trochozoa 325 16 Molluscs
338
17 Annelids
369
18 Smaller Ecdysozoans
Glossary G-1 Index I-1
391
19 Trilobites, Chelicerates, and Myriapods 409
iii
TABLE OF CONTENTS
About the Authors ix Preface x
PART ONE
3.3 Mitosis and Cell Division 51 Summary 54
CHAPTER 4 Cellular Metabolism 57 4.1 Energy and the Laws of Thermodynamics 58 4.2 The Role of Enzymes 59 4.3 Enzyme Regulation 61 4.4 Chemical Energy Transfer by ATP 62 4.5 Cellular Respiration 63 4.6 Metabolism of Lipids 69 4.7 Metabolism of Proteins 70 Summary 71
PART T WO Introduction to Living Animals CHAPTER 1 Life: Biological Principles and the Science of Zoology 1 1.1 Fundamental Properties of Life 2 1.2 Zoology as a Part of Biology 9 1.3 Principles of Science 9 1.4 Theories of Evolution and Heredity 13 Summary 18
CHAPTER 2 The Origin and Chemistry of Life 20 2.1 Water and Life 21 2.2 Macromolecules 23 2.3 Chemical Evolution 26 2.4 Origin of Living Systems 29 2.5 Precambrian Life 31 Summary 34
CHAPTER 3 Cells as Units of Life 36 3.1 Cell Concept 37 3.2 Organization of Cells 39
iv
Continuity and Evolution of Animal Life CHAPTER 5 Genetics: A Review 73 5.1 Mendel’s Investigations 74 5.2 Chromosomal Basis of Inheritance 74 5.3 Mendelian Laws of Inheritance 79 5.4 Gene Theory 88 5.5 Storage and Transfer of Genetic Information 88 5.6 Gene Mutations 98 5.7 Molecular Genetics of Cancer 99 Summary 99
Table of Contents
CHAPTER 6
CHAPTER 9
Organic Evolution 103
Architectural Pattern of an Animal 187
6.1 Origins of Darwinian Evolutionary Theory 104 6.2 Darwinian Evolutionary Theory: The Evidence 107 6.3 Revisions of Darwin’s Theory 125 6.4 Microevolution: Genetic Variation and Change Within Species 126 6.5 Macroevolution: Major Evolutionary Events 132 Summary 134
9.1 Hierarchical Organization of Animal Complexity 188 9.2 Animal Body Plans 189 9.3 Components of Animal Bodies 193 9.4 Complexity and Body Size 198 Summary 199
CHAPTER 7 The Reproductive Process 137
CHAPTER 10 Taxonomy and Phylogeny of Animals 201
7.1 Nature of the Reproductive Process 138 7.2 The Origin and Maturation of Germ Cells 142 7.3 Reproductive Patterns 146 7.4 Structure of Reproductive Systems 147 7.5 Endocrine Events That Orchestrate Reproduction 149 Summary 156
10.1 Linnaeus and Taxonomy 202 10.2 Species 204 10.3 Taxonomic Characters and Phylogenetic Reconstruction 209 10.4 Theories of Taxonomy 211 10.5 Major Divisions of Life 217 10.6 Major Subdivisions of the Animal Kingdom 217 Summary 218
CHAPTER 8
CHAPTER 11
Principles of Development 158
Unicellular Eukaryotes 220
8.1 Early Concepts: Preformation Versus Epigenesis 159 8.2 Fertilization 160 8.3 Cleavage and Early Development 163 8.4 An Overview of Development Following Cleavage 165 8.5 Mechanisms of Development 167 8.6 Gene Expression During Development 170 8.7 Developmental Patterns in Animals 173 8.8 Evolutionary Developmental Biology 178 8.9 Vertebrate Development 178 8.10 Development of Systems and Organs 181 Summary 185
11.1 Naming and Identifying Unicellular Eukaryotic Taxa 221 11.2 Form and Function 225 11.3 Major Unicellular Eukaryotic Taxa 231 11.4 Phylogeny and Adaptive Diversification 245 Summary 248
PART THREE
CHAPTER 12 Sponges and Placozoans 250 12.1 Origin of Animals 251 12.2 Phylum Porifera: Sponges 251 12.3 Phylum Placozoa 262 Summary 263
CHAPTER 13 Cnidarians and Ctenophores 264 13.1 Phylum Cnidaria 265 13.2 Phylum Ctenophora 287 13.3 Phylogeny and Adaptive Diversification 290 Summary 292
CHAPTER 14 Xenacoelomorpha, Platyzoa, and Mesozoa 294
Diversity of Animal Life
14.1 Phylum Xenacoelomorpha 295 14.2 Clades Within Protostomia 297 14.3 Phylum Platyhelminthes 297 14.4 Phylum Gastrotricha 312 14.5 Clade Gnathifera 313
v
vi
Table of Contents
14.6 Phylum Gnathostomulida 314 14.7 Phylum Micrognathozoa 315 14.8 Phylum Rotifera 316 14.9 Phylum Acanthocephala 318 14.10 Phylum Mesozoa 320 14.11 Phylogeny 320 Summary 322
CHAPTER 15 Polyzoa and Trochozoa 325 15.1 Clade Polyzoa 327 15.2 Phylum Cycliophora 327 15.3 Phylum Entoprocta 327 15.4 Phylum Ectoprocta (Bryozoa) 328 15.5 Clade Trochozoa 331 15.6 Clade Brachiozoa 331 15.7 Phylum Brachiopoda 331 15.8 Phylum Phoronida 332 15.9 Phylum Nemertea (Rhynchocoela) 333 15.10 Phylogeny 335 Summary 336
CHAPTER 16 Molluscs 338 16.1 Molluscs 339 16.2 Form and Function 341 16.3 Classes of Molluscs 344 16.4 Phylogeny and Adaptive Diversification 364 Summary 367
CHAPTER 17 Annelids 369 17.1 Phylum Annelida, Including Pogonophorans (Siboglinids), Sipunculans, and Echiurans 371 17.2 Errantia 374 17.3 Sedentaria 376 17.4 Clade Clitellata 381 17.5 Evolutionary Significance of a Coelom and Metamerism 388 17.6 Phylogeny and Adaptive Diversification 389 Summary 389
CHAPTER 18 Smaller Ecdysozoans 391 18.1 Phylum Nematoda: Roundworms 392 18.2 Phylum Nematomorpha 400 18.3 Phylum Loricifera 401 18.4 Phylum Kinorhyncha 401 18.5 Phylum Priapulida 402 18.6 Clade Panarthropoda 403 18.7 Phylum Onychophora 403
18.8 Phylum Tardigrada 404 18.9 Phylogeny and Adaptive Diversification 406 Summary 407
CHAPTER 19 Trilobites, Chelicerates, and Myriapods 409 19.1 Phylum Arthropoda 410 19.2 Subphylum Trilobita 413 19.3 Subphylum Chelicerata 414 19.4 Subphylum Myriapoda 422 19.5 Phylogeny and Adaptive Diversification 425 Summary 426
CHAPTER 20 Crustaceans 428 20.1 Subphylum Crustacea 430 20.2 A Brief Survey of Crustaceans 438 20.3 Phylogeny and Adaptive Diversification 447 Summary 448
CHAPTER 21 Hexapods 450 21.1 Classes Entognatha and Insecta 451 21.2 External Form and Function 451 21.3 Internal Form and Function 457 21.4 Metamorphosis and Growth 464 21.5 Behavior and Defense 466 21.6 Insects and Human Welfare 470 21.7 Phylogeny and Adaptive Diversification 476 Summary 477
CHAPTER 22 Chaetognaths, Echinoderms, and Hemichordates 480 22.1 Phylum Chaetognatha 482 22.2 Form and Function 482 22.3 Clade Ambulacraria 483 22.4 Phylum Echinodermata 483 22.5 Phylogeny and Adaptive Diversification of Echinoderms 499 22.6 Phylum Hemichordata 500 22.7 Phylogeny and Adaptive Diversification of Hemichordates 503 Summary 504
CHAPTER 23 Chordates 506 23.1 Ancestry and Evolution of the Chordates 507 23.2 Five Chordate Characteristics 510 23.3 Subphylum Urochordata 512 23.4 Subphylum Cephalochordata 514
23.5 Subphylum Vertebrata 515 Summary 522
Table of Contents
vii
PART FOUR
CHAPTER 24 Fishes 524 24.1 Ancestry and Relationships of Major Groups of Fishes 525 24.2 Living Jawless Fishes 525 24.3 Chondrichthyes: Cartilaginous Fishes 531 24.4 Osteichthyes: Bony Fishes and Tetrapods 535 24.5 Structural and Functional Adaptations of Fishes 539 Summary 549
CHAPTER 25 Early Tetrapods and Modern Amphibians 552 25.1 Devonian Origin of Tetrapods 553 25.2 Temnospondyls and Modern Amphibians 555 25.3 Caecilians: Order Gymnophiona (Apoda) 557 25.4 Salamanders: Order Urodela (Caudata) 557 25.5 Frogs and Toads: Order Anura (Salientia) 560 Summary 569
CHAPTER 26 Amniote Origins and Nonavian Reptiles 571 26.1 Origin and Early Evolution of Amniotes 572 26.2 Characteristics and Natural History of Reptilian Groups 577 Summary 591
CHAPTER 27 Birds 593 27.1 Origin and Relationships 594 27.2 Structural and Functional Adaptations for Flight 597 27.3 Flight 605 27.4 Migration and Navigation 608 27.5 Social Behavior and Reproduction 610 27.6 Bird Populations and Their Conservation 613 Summary 617
CHAPTER 28 Mammals 619 28.1 Origin and Evolution of Mammals 620 28.2 Structural and Functional Adaptations of Mammals 623 28.3 Humans and Mammals 637 28.4 Human Evolution 638 Summary 646
Activity of Life CHAPTER 29 Support, Protection, and Movement 648 29.1 Integument 649 29.2 Skeletal Systems 652 29.3 Animal Movement 658 Summary 667
CHAPTER 30 Homeostasis: Osmotic Regulation, Excretion, and Temperature Regulation 669 30.1 Water and Osmotic Regulation 670 30.2 Invertebrate Excretory Structures 674 30.3 Vertebrate Kidney 676 30.4 Temperature Regulation 682 Summary 686
CHAPTER 31 Homeostasis: Internal Fluids and Respiration 689 31.1 Internal Fluid Environment 690 31.2 Composition of Blood 691 31.3 Circulation 693 31.4 Respiration 701 Summary 709
CHAPTER 32 Digestion and Nutrition 711 32.1 Feeding Mechanisms 712 32.2 Digestion 715 32.3 Organization and Regional Function of Alimentary Canals 717 32.4 Regulation of Food Intake 722
viii
Table of Contents
32.5 Nutritional Requirements 724 Summary 727
PART FIVE
CHAPTER 33 Nervous Coordination: Nervous System and Sense Organs 729 33.1 Neurons: Functional Units of Nervous Systems 730 33.2 Synapses: Junctions Between Nerves 734 33.3 Evolution of Nervous Systems 736 33.4 Sense Organs 742 Summary 753
CHAPTER 34 Chemical Coordination: Endocrine System 755 34.1 Mechanisms of Hormone Action 756 34.2 Invertebrate Hormones 758 34.3 Vertebrate Endocrine Glands and Hormones 760 Summary 770
CHAPTER 35 Immunity 773 35.1 Susceptibility and Resistance 774 35.2 Innate Defense Mechanisms 774 35.3 Immunity in Invertebrates 778 35.4 Acquired Immune Response in Vertebrates 778 35.5 Blood Group Antigens 784 Summary 785
CHAPTER 36 Animal Behavior 787 36.1 Describing Behavior: Principles of Classical Ethology 789 36.2 Control of Behavior 790 36.3 Social Behavior 794 Summary 803
Animals and Their Environments CHAPTER 37 Animal Distributions 806 37.1 Principles of Historical Biogeography 807 37.2 Distribution of Life on Earth 813 Summary 824
CHAPTER 38 Animal Ecology 826 38.1 The Hierarchy of Ecology 827 38.2 Populations 828 38.3 Community Ecology 832 38.4 Ecosystems 836 38.5 Extinction and Biodiversity 840 Summary 843 Glossary G-1 Index I-1
ABOUT THE AUTHORS
CLEVELAND P. HICKMAN, JR. Cleveland P. Hickman, Jr., Professor Emeritus of Biology at W ashington and Lee University in Lexington, Virginia, has taught zoology and animal physiology for more than 30 years. He received his Ph.D. in comparative physiology from the University of British Columbia, Vancouver, B.C., in 1958 and taught animal physiology at the University of Alberta before moving to Washington and Lee University in 1967. He has published numerous articles and research papers in fish physiology, in addition to co-authoring these highly successful texts: Integrated Principles of Zoology, Biology of Animals, Animal Diversity, Laboratory Studies in Animal Diversity, and Laboratory Studies in Integrated Principles of Zoology. Over the years Dr. Hickman has led many field trips to the Galápagos Islands. His research is on intertidal zonation and marine invertebrate systematics in the Galápagos. He has published four field guides in the Galápagos Marine Life Series for the identification of echinoderms, marine molluscs, corals, and marine crustaceans. His interests include scuba diving, woodworking, and participating in chamber music ensembles. Dr. Hickman can be contacted at: [email protected]
SUSAN KEEN Susan Keen is a Senior Lecturer in the College of Biological Sciences, University of California at Davis. She was Associate Dean for Undergraduate Academic Programs from 2011 to 2018. She received her Ph.D. in zoology from the University of California–Davis, following a M.Sc. from the University of Michigan at Ann Arbor. She is a native of Canada and obtained her undergraduate education at the University of British Columbia in Vancouver. Dr. Keen is an invertebrate zoologist fascinated with sea jelly life histories, in particular those where both asexual and sexual phases of organisms are present, as in most jellies. Other research has included work on sessile marine invertebrate communities, spider populations,
Andean potato evolution, and most recently how students learn. Dr. Keen has been teaching evolution and animal diversity within the Introductory Biology series for 25 years. She enjoys all facets of the teaching process, from lectures and discussions to the design of effective laboratory exercises. In addition to her work with introductory biology, she works with an animator to create self-guided teaching modules on animal development. She attended the National Academies Summer Institute on Undergraduate Education in Biology, and was a National Academies Education Fellow in the Life Sciences for 2005–2006. Her interests include horseback riding, gardening, travel, and mystery novels. Dr. Keen can be contacted at: [email protected]
DAVID J. EISENHOUR David J. Eisenhour is professor of biology at Morehead State University in Morehead, Kentucky. He received his Ph.D. in zoology from Southern Illinois University, Carbondale. He teaches courses in environmental science, human anatomy, mammalogy, general zoology, comparative anatomy, ichthyology, and vertebrate zoology. David has an active research program that focuses on systematics, conservation biology, and natural history of North American freshwater fishes. He has a particular interest in the diversity of Kentucky’s fishes and is writing a book about that subject. He and his students have authored several publications. David serves as an academic advisor to prepharmacy students. His interests include fishing, landscaping, softball, hiking, and entertaining his three children, who, along with his wife, are enthusiastic participants in fieldwork. Dr. Eisenhour can be contacted at: [email protected]
ALLAN LARSON Allan Larson is a professor at Washington University, St. Louis, MO. He received his Ph.D. in genetics at the University of California–Berkeley, following B.S. and M.S. degrees in zoology from the University
of Maryland. His fields of specialization include evolutionary biology, molecular population genetics and systematics, and amphibian systematics. He teaches courses in introductory genetics, zoology, macroevolution, molecular evolution, and the history of evolutionary theory, and has organized and taught a special course in evolutionary biology for high-school teachers. Dr. Larson’s students have participated in zoological field studies around the world, including projects in Africa, Asia, Australia, Madagascar, North America, South America, the Indo-Pacific Ocean, and the Caribbean Islands. Dr. Larson has authored numerous scientific publications, and has edited for the journals The American Naturalist, Evolution, Journal of Experimental Zoology, Molecular Phylogenetics and Evolution, and Systematic Biology. Dr. Larson serves as an academic advisor to undergraduate students and directs undergraduate studies in biology at Washington University. Dr. Larson can be contacted at: [email protected]
HELEN I’ANSON Helen I’Anson, a native of England, is the John T. Perry Professor of Biology and Research Science at Washington and Lee University in Lexington, Virginia. She received her Ph.D. in physiology at the University of Kentucky, Lexington, and postdoctoral training at the University of Michigan, Ann Arbor. She teaches courses in animal physiology, microanatomy, neuroendocrinology, general biology, and reproductive physiology. She has an active research program that focuses on the mechanisms regulating the development of childhood obesity. Currently she is studying the role of snacking from weaning to adulthood in the onset of childhood obesity. In 2019 she was awarded the Outstanding Faculty Award for teaching, research, and service by the State Council of Higher Education for Virginia. Her interests include gardening, hiking, fishing, aromatherapy, music, and participating in choral ensembles. Dr. I’Anson can be contacted at: [email protected] ix
P RE FA C E
I
ntegrated Principles of Zoology continues to be the leading text for the introductory zoology course. With the eighteenth edition, the authors bring a wealth of real experience as they describe the diversity of animal life and the fascinating adaptations that enable animals to inhabit so many ecological niches. The overall organization of this text has proven to work well to help students understand the content. Distinctive features, especially the emphasis on principles of evolution and zoological science, have been strengthened. To aid in student learning, several pedagogical features have been retained: opening chapter prologues drawn from the chapter’s theme; chapter summaries and review questions to aid in comprehension and study; concise and visually appealing illustrations; chapter notes and essays that offer interesting sidelights to the narrative; literature citations; and an extensive glossary providing pronunciations, derivations, and definitions of terms used in the text.
NEW TO THE EIGHTEENTH EDITION Starting with this edition, a list of Learning Objectives opens each chapter. These objectives are organized according to the chapter’s main sections. Chapter summaries, many of which are expanded in content, are restructured in tabular form to list the main highlights of each section of the chapter. This correspondence between the learning objectives, chapter sections, and summary should help students to organize the main lessons of each chapter. Our extensive cross-referencing of material among the different parts of the book now uses section numbers, with live links available in electronic versions of the text. Notes and essays separate from the main text are now numbered for reference and organized according to six Key Themes: (1) Adaptation and Physiology, (2) Ecology, (3) Evolution, (4) Genetics and Development, (5) Human Connections, and (6) Science Explained. Adaptation and Physiology connects the proximate and ultimate causes underlying organismal functioning. Ecology addresses the interactions of animal populations with their environments, including factors that influence their geographic distribution and abundance. Evolution highlights common descent of animal species and the historical processes that modify organismal characteristics in natural populations. Genetics and Development covers the mechanisms of heredity and the ways in which an organism uses genetic information to progress from a zygote ultimately to an adult animal. Human Connections highlights ways in which zoological findings influence human welfare, including medical applications and maintaining environmental health, or how human activities have affected animal species. Science Explained covers scientific methodology and the history of scientific discovery. In addition to these organizational revisions, we have replaced many photographs and diagrams throughout the book to improve clarity and vibrancy. x
We have updated geological periods on our phylogenetic trees throughout the book; for example, the former Tertiary period is replaced with the Paleogene and Neogene periods. We highlight some major revisions in order by the book’s five major parts.
Part One, Introduction to Living Animals In Chapter 1, we introduce the microbiome as an important characteristic of animal life. It is often overlooked that animals typically harbor thousands of species of bacteria and archaea, primarily in the gut. These species typically exist in a harmless symbiosis with their animal hosts, with fewer than 100 species of bacteria being sources of infectious disease. As molecular genetic methods for identifying an animal’s microbiome improve, this dimension is becoming an increasingly important part of zoology. In Chapter 2, water as a solvent for respiratory gases is added to its important properties for supporting life. The introduction to proteins is made more specific in stating that a typical protein contains hundreds to thousands of amino acids of 20 standard kinds. Prions are described in a Key Theme essay. Atmospheric carbon dioxide is updated to 0.04%. The name Archaebacteria is updated to Archaea. In discussing endosymbiotic theory, cyanobacterial invaginations are introduced as a corresponding feature between cyanobacteria and plastids. Chapter 3 includes more explicit labeling of some figures and color coordination among figures that incorporate the plasma membrane.
Part Two, Continuity and Evolution of Animal Life In Chapter 5, single nucleotide polymorphism (SNP) is made a major concept and given a glossary entry. Key Themes include titles on chromosomes and the cell cycle, genic cloning, genome size, and junk or selfish DNA, and radiation and chemical mutagens. The chapter summary is expanded to cover more details of the chapter. Chapter 6 includes new material on neutral alleles and species selection in connection with the learning objectives. August Weismann’s role in neo-Darwinian theory is elaborated. In Chapter 7, parthenogenesis has been moved from the section on sexual reproduction and presented instead as a separate condition intermediate between sexual and asexual reproduction. Chapter 8 includes a new short discussion of coelom function. Chapter 9 includes new material on the organizational level of flatworms, the status of the spongocoel as a gut cavity in sponges, junctional complexes found between epithelial cells, and properties of oblique striated muscle.
Preface xi
Part Three, Diversity of Animal Life In Chapter 10, new information is added on the roles of the holotype and paratypes in taxonomy, with new glossary definitions of these terms. Contents of the major subdivisions of animal life are updated to reflect new phylogenetic hypotheses. Chapter 11 includes three new glossary terms: ancoracyst, mixotrophy, and trogocytosis, plus a revised description of red tide in a Key Theme. New descriptions of clades Holomyota and Holozoa (within Opisthokonta) appear with an associated revised Figure 11.36. Chapter 12 has a new opening essay and a revised section on the origin of animals, with bioactive compounds now a Key Theme. Class Myxozoa has been added to Chapter 13 (full description) and to the associated cladogram in Figure 13.2. Phylum Xenacoelomorpha is now included in Chapter 14: Xenoturbellida, previously in Chapter 22, is now considered a protostome related to acoelomorph flatworms and not a deuterostome, so it has been allied with Acoelomorpha as Xenacoelomorpha. The figure from Chapter 22 is now Figure 14.2. The phylogeny section explains the revision of the group. Chapter 14 also contains a revised Key Theme on schistosomiasis. In Chapter 15, clade Kryptrochozoa has been removed in favor of the more inclusive group Trochozoa, with an associated change in the cladogram, Figure 15.1. In Chapter 17, Phylum Sipuncula has been demoted and placed within Phylum Annelida; the cladogram in Figure 17.1 is revised to accommodate this change. Several new section numbers added to Chapter 17 make it easier to follow major topics. Chapter 18 contains a new Key Theme on cryptobiosis and its applications to storing blood. Chapter 21 contains a new Key Theme on Varroa mites and bee colony collapse disorder. The opening essay has updated statistics on locust outbreaks. Several new section numbers were added here to make it easier to follow major topics. Chapter 22 includes a revised Key Theme essay on Crown-of-Thorns sea-star outbreaks and one on Diadema sea urchin outbreaks. Figure 22.3 is revised to include new echinoderm fossil groups, which are discussed in the echinoderm phylogeny section. Several new section numbers make it easier to follow major topics. Chapter 23 features consolidation of material on early chordate evolution into a single section, and improved art for the cladogram, chordate characters, and vertebrate innovations. New text is added to explain problems with using Linnean taxonomy in chordates, and why an unranked system is favored for higher chordate taxonomy. This material applies as well to the other vertebrate chapters. Taxonomy and species numbers are updated. The biology and ecology of amphioxus get increased treatment to match that of sea squirts and to introduce traits discussed in the vertebrate chapters. New material on early vertebrate evolution includes the early chordate Metaspriggina. In Chapter 24, the introduction is modestly revised to outline the contents of the chapter. A single table now consolidates characteristics of the four major fish groups, allowing easy comparison among groups. Teleost suction feeding is expanded to emphasize its importance and to contrast it with feeding in terrestrial vertebrates. Shark reproduction is rewritten to emphasize continuity of reproductive mechanisms. Chapter 25 is reorganized into a greater number of sections with an expanded summary and some new information on temnospondyls as a source of modern amphibians. Numbers of species are updated to represent recent discoveries of cryptic species in many taxa. The
introduction to Chapter 26 is reorganized to outline the chapter’s contents. Changes to the section on amniote adaptations include revised coverage of respiration, circulation, and special sensory systems. Some material on limb positions was moved to the mammal chapter. New text on squamate reproduction complements coverage for turtles, crocodilians, and tuataras. Material on snake venom is rewritten for clarity. The section on dinosaur systematics is updated. Chapter 27 includes a new paragraph on early bird evolution in the Cretaceous and Paleogene, and an extensive rewrite on dinosaur ancestry of birds. We revised the sections on fluid dynamics of avian flight and the respiratory cycle. An expanded boxed essay now covers lead poisoning of condors. We reduce usage of the somewhat awkward term “nonavian reptile” for the traditional Class Reptilia. Chapter 28 features a substantial new section on early mammalian evolution and on the megafauna of the Pleistocene. Updates include mammalian dentition and feeding, migration of bats, and decline of caribou populations in North America. Coverage of mammalian population ecology was reduced to avoid redundancy with Chapter 38. Human evolution is revised to include new research on Australopithecus, Homo heidelbergensis, and Denisovans.
Part Four, Activity of Life Chapter 30 includes new information on how mixing of freshwater and seawater at the ebbing of tides produces physiological challenges for aquatic animals. Oncotic pressure (colloid osmotic pressure) is added as a force opposing blood hydrostatic pressure and as a factor that produces edema in blood or interstitial fluid. New emphasis is placed on the relationship between surface area and volume in mammals’ ability to withstand cold environmental temperatures. Chapter 31 includes a new Key Theme on hypertension and associated kidney disease. Chapter 32 adds breakdown of nucleic acids to the section on digestion. Chapter 33 features redrawn Figures 33.7 and 33.8, with channels shown as opening when the neurotransmitter binds. In Chapter 34 a new Key Theme lists the heart as an endocrine organ producing atrial natriuritic peptide. New information is added on the roles of oxytocin and vasopressin in social behavior and the use of oxytocin as a treatment for autism spectrum disorders. In Chapter 36, polyandry is now illustrated with the example of Galápagos hawks.
Part Five, Animals and Their Environments In Chapter 37, a new Key Theme is added on the demand for fresh water and how global warming challenges our supply of fresh water. Information on the Peruvian anchoveta fishery is updated. Chapter 38 is restructured into more sections to coordinate the learning objectives and chapter summary. The growth of human populations and the taxonomic status of Galápagos finches are updated.
TEACHING AND LEARNING AIDS To help students in vocabulary development, key words are boldfaced and derivations of technical and zoological terms are provided, along with generic names of animals where they first appear in the text. In this way students gradually become familiar with the
xii
Preface
more common roots that form many technical terms. An extensive glossary provides pronunciation, derivation, and definition for many terms, including new ones added to the glossary or existing ones rewritten for this edition. A distinctive feature of this text is a prologue for each chapter that highlights a theme or fact relating to the chapter. Some prologues present biological, particularly evolutionary, principles; those in Part Three on animal diversity illuminate distinguishing characteristics of the group presented in the chapter. Again, William C. Ober and Claire W. Ober have strengthened the art program for this text with many new full-color paintings that replace older art, or that illustrate new material. Bill’s artistic skills, knowledge of biology, and experience gained from an earlier career as a practicing physician have enriched this text through 10 of its editions. Claire practiced pediatric and obstetric nursing before turning to scientific illustration as a full-time career. Texts illustrated by Bill and Claire have received national recognition and won awards from the Association of Medical Illustrators, American Institute of Graphic Arts, Chicago Book Clinic, Printing Industries of America, and Bookbuilders West. They are also recipients of the Art Directors Award.
For the Zoology Lab Laboratory Studies in Integrated Principles of Zoology by Cleveland Hickman, Jr., Susan Keen, and Lee B. Kats Now in its eighteenth edition, this lab manual was written to accompany Integrated Principles of Zoology, and can be easily adapted to fit a variety of course plans.
ACKNOWLEDGMENTS We have received suggestions from faculty and students throughout the country. This is vital feedback that we rely on with each edition. Each person who has offered comments and suggestions has our thanks. The efforts of many people are needed to develop and improve a textbook. Among these people are the reviewers and consultants who point out areas of concern, cite areas of strength, and make recommendations for change. The following reviewers helped review the seventeenth edition to help with development of the eighteenth: Richard S. Grippo, Ph.D., Arkansas State University Melissa Gutierrez, MS, The University of Southern Mississippi Patrick J. Lewis, Ph.D., Sam Houston State University Matthew Nusnbaum, PhD, Georgia State University Amy Reber, Ph.D., Georgia State University Natalie Reynolds, MsEd, Carl Albert State College Rita A. Thrasher, MS, Pensacola State College Travis J. Vail, MS, Golden West College The authors express their appreciation to the editors and support staff at McGraw-Hill Higher Education who made this project possible. Special thanks are due to Michael Ivanov, Senior Portfolio Manager; Michelle Flomenhoft, Senior Product Developer; Erin DeHeck, Product Developer; Kelly Brown; Senior Marketing Manager; Becca Gill, Content Project Manager; Ann Courtney, Senior Content Project Manager; Brent dela Cruz, Lead Content Project Manager; Jessica Cuevas, Designer; and Laura Fuller, Senior Buyer. We are indebted to them for their talents and dedication. Although we make every effort to bring to you an errorfree text, errors of many kinds inevitably find their way into a textbook of this scope and complexity. We will be grateful to readers who have comments or suggestions concerning content and send their remarks to your McGraw-Hill sales representative. To find your McGraw-Hill representative, go to www.mheducation.com and click “Get Support,” select “Higher Ed” and then click the “Get Started” button under the “Find Your Sales Rep” section. Cleveland P. Hickman, Jr. Susan Keen David J. Eisenhour Allan Larson Helen I’Anson
Preface xiii
CONNECT FOR INTEGRATED PRINCIPLES OF ZOOLOGY, 18e McGraw-Hill Connect for Integrated Principles of Zoology, 18e will include: ∙∙
Question Bank ∙∙ Multiple Choice questions ∙∙ Check All That Apply questions ∙∙ Interactive exercises, such as matching, classification, labeling, and sequencing activities
∙∙
Testbank ∙∙ Multiple Choice questions ∙∙ Fill-in-the-Blank questions ∙∙ Short Answer questions
INSTRUCTORS’ RESOURCES You can rely on the following materials to help you and your students work through the material in the book; all are available in the Instructor Resources under the library tab in Connect (available only to instructors who are logged in to Connect). Supplement
Features
Instructor’s Manual
∙ Chapter Outlines ∙ Lesson Plans ∙ Advanced Class Questions ∙ Answer Key for End-of-Chapter questions (posted separately)
Presentation Tools
∙ Accessible Lecture PowerPoints ∙ Image PowerPoints ∙ Baseart Images ∙ Labeled Art Images ∙ Animations
Electronic Testbank
∙ Computerized and Connect ∙ Word Version ∙ Questions tagged for Learning Outcomes, Level of Bloom’s Taxonomy, Section, and Topic
Professional Resources
Links to websites of interest
Chapter References
Listing of Selected References for each chapter (previously listed in the book)
Endsheets
∙ Cladogram ∙ Timeline of Events
Conversion Guides
Spreadsheets detailing changes in each chapter between 17e and 18e
The Structure of Matter
PDF of Structure of Matter Appendix that appeared in earlier editions of the book
Additional Labs
∙ Lab Manual References (as noted in Appendix A of Lab Manual) ∙ PDFs of Additional Labs that appeared in earlier editions of the lab manual
Want to learn more about this product? Attend one of our online webinars. To learn more about them, please contact your McGraw-Hill sales representative. To find your McGraw-Hill representative, go to www.mheducation.com and click “Get Support,” select “Higher Ed” and then click the “Get Started” button under the “Find Your Sales Rep” section. Need Help? Contact the McGraw-Hill Education Customer Experience Group (CXG) Visit the CXG website at www.mhhe.com/support. Browse our frequently asked questions (FAQs) and product documentation and/or contact a CXG representative.
®
FOR INSTRUCTORS You’re in the driver’s seat. Want to build your own course? No problem. Prefer to use our turnkey, prebuilt course? Easy. Want to make changes throughout the semester? Sure. And you’ll save time with Connect’s auto-grading too.
65%
Less Time Grading
They’ll thank you for it. Adaptive study resources like SmartBook® 2.0 help your students be better prepared in less time. You can transform your class time from dull definitions to dynamic debates. Find out more about the powerful personalized learning experience available in SmartBook 2.0 at www.mheducation.com/highered /connect/smartbook Laptop: McGraw-Hill Education
Make it simple, make it affordable.
Solutions for your challenges.
Connect makes it easy with seamless integration using any of the major Learning Management Systems— Blackboard®, Canvas, and D2L, among others—to let you organize your course in one convenient location. Give your students access to digital materials at a discount with our inclusive access program. Ask your McGraw-Hill representative for more information.
A product isn’t a solution. Real solutions are affordable, reliable, and come with training and ongoing support when you need it and how you want it. Our Customer Experience Group can also help you troubleshoot tech problems— although Connect’s 99% uptime means you might not need to call them. See for yourself at status. mheducation.com
Padlock: Jobalou/Getty Images
Checkmark: Jobalou/Getty Images
FOR STUDENTS Effective, efficient studying. Connect helps you be more productive with your study time and get better grades using tools like SmartBook 2.0, which highlights key concepts and creates a personalized study plan. Connect sets you up for success, so you walk into class with confidence and walk out with better grades.
Study anytime, anywhere. Download the free ReadAnywhere app and access your online eBook or SmartBook 2.0 assignments when it’s convenient, even if you’re offline. And since the app automatically syncs with your eBook and SmartBook 2.0 assignments in Connect, all of your work is available every time you open it. Find out more at www.mheducation.com/readanywhere
“I really liked this app—it made it easy to study when you don't have your textbook in front of you.” - Jordan Cunningham, Eastern Washington University
No surprises. The Connect Calendar and Reports tools keep you on track with the work you need to get done and your assignment scores. Life gets busy; Connect tools help you keep learning through it all. Calendar: owattaphotos/Getty Images
Learning for everyone. McGraw-Hill works directly with Acc