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Fourth Edition

Richard C. Brusca, Gonzalo Giribet, and Wendy Moore

20 December 2022

ISBN: 9780197554418

1104 pages

Price: £128.99

An invaluable text and reference for both students and scholars alike, this is the most complete, authoritative, and visually engaging guide to the field of invertebrate biology. If you are a lecturer interested in adopting this title for your course, please contact your local campus representative to arrange a local price.



An invaluable text and reference for both students and scholars alike, Invertebrates is the most complete, authoritative, and visually engaging guide to the field of invertebrate biology.

  • Retains the popular aspects of previous editions, but is substantially shorter in length
  • Detailed classifications and taxonomic synopses within each phylum are provided. These are to be used as a reference to look up taxonomic names, understand the traits that distinguish groups, or get an overall sense of the scope of the higher taxa in a phylum.
  • Invertebrates is the only college textbook on the subject that is in full color.
  • Introductory chapters introduce students to systematics and phylogenetics, reviewing animal body plans, metazoan development, and other fundamentals of zoology.

About the Author(s)

Richard C. Brusca, University of Arizona, Gonzalo Giribet, Harvard University, and Wendy Moore, University of Arizona

Richard C. Brusca, Ph.D., University of Arizona, is Executive Director Emeritus of the Arizona-Sonora Desert Museum and a Research Scientist in the Department of Ecology and Evolutionary Biology at the University of Arizona. Dr. Brusca is the author of over 200 research publications and 15 books, and the recipient of more than 100 research grants from the National Science Foundation, NOAA, and the National Geographic Society, and many others. He is a Fellow of the American Association for the Advancement of Science, the Linnean Society of London, and the California Academy of Sciences. His research interests include invertebrate zoology, the natural history of the Sonoran Desert and Gulf of California, and ecology of Arizona's Madrean Sky Islands.

Gonzalo Giribet, Ph.D., University of Barcelona, is Alexander Agassiz Professor of Zoology, Curator of Invertebrates and Director of the Museum of Comparative Zoology, Harvard, and a Professor in the Department of Organismic and Evolutionary Biology. In addition, he holds an Doctor Honoris Causa from the University of Copenhagen, and he an Honorary Research Fellow at The Natural History Museum, London, a Fellow of the California Academy of Sciences, and a 2016 Fellow of the John Simon Guggenheim Memorial Foundation. He has published more than 400 scientific articles and book chapters, and has named more than 100 new species, genera, and families of invertebrates. His research is broadly interested in the origins and maintenance of animal diversity, which he explores through studying the evolution, systematics, and biogeography of invertebrate animals.

Wendy Moore, Ph.D., University of Arizona, is Associate Professor in the Department of Entomology at the University of Arizona and Curator of the University of Arizona Insect Collection. Her long-term research interest is the evolution of diversity, particularly of invertebrates and especially of insects. She is a systematist who uses molecular-based phylogenetics to understand the evolution of morphology and how biotic, climatic, and tectonic events may have influenced the timing and patterns of diversification. She is also committed to natural history collections care and enhancement, and the use of bioinformatics to make collections-based data widely available to diverse user communities.

Table of Contents

    Chapter 1
    Keeping Track of Life
    Prokaryotes and Eukaryotes
    Where Did Invertebrates Come From?
    Where Do Invertebrates Live?
    Changing Views of Invertebrate Phylogeny
    A Final Introductory Message to the Reader
    Chapter 2
    Systematics, Phylogeny, and Classification
    Phylogeny, Monophyly, Paraphyly, and Polyphyly
    Apomorphy and Plesiomorphy
    Challenges of Phylogenetic Inference
    Constructing Phylogenies
    Biological Classification
    Chapter 3
    Introduction to the Animal Kingdom: Animal Architecture and Body Plans
    Body Symmetry
    Cellularity, Body Size, Germ Layers, and Body Cavities
    Locomotion and Support
    Cilia and Flagella
    Feeding and Digestion
    Excretion and Osmoregulation
    Circulation and Gas Exchange
    Nervous Systems and Sense Organs
    Nervous Systems and Body Plans
    Hormones and Pheromones
    Chapter 4
    Introduction to the Animal Kingdom: Development, Life Histories, and Origin
    Evolutionary Developmental Biology: Evo-Devo
    Eggs and Embryos
    Life Cycles: Sequences and Strategies
    The Relationships Between Ontogeny and Phylogeny
    The Origin of the Metazoa
    Chapter 5
    Phylum Porifera: The Sponges
    Taxonomic History and Classification
    The Poriferan Body Plan
    Some Additional Aspects of Sponge Biology
    Poriferan Phylogeny
    Chapter 6
    Two Enigmatic Phyla: Placozoa and Ctenophora (The Comb Jellies)
    Phylum Ctenophora
    Taxonomic History and Classification
    The Ctenophoran Body Plan
    Ctenophoran Phylogeny
    Chapter 7
    Phylum Cnidaria: Anemones, Corals, Jellyfish, and Their Kin
    Taxonomic History and Classification
    The Cnidarian Body Plan
    Cnidarian Evolutionary History
    Chapter 8
    A Brief introduction to the Bilateria and Its Major Clades
    Deuterostomes and Protostomes
    Chapter 9
    Phylum Xenacoelomorpha. Basal Bilaterians
    The Basal Bilaterian
    Phylum Xenacoelomorpha
    Subphylum Acoelomorpha
    Class Acoela
    The Acoel Body Plan
    Class Nemertodermatida
    The Nemertodermatid Body Plan
    Subphylum Xenoturbellida
    The Xenoturbellid Body Plan
    Chapter 10
    Protostomia, Spiralia, and the Phylum Dicyemida
    Spiralia and Ecdysozoa
    The Phylum Dicyemida (= Rhombozoa)
    Anatomy and Biology of Dicyemidans
    Life Cycles
    Chapter 11
    Gnathifera. The Phyla Gnathostomulida, Rotifera (including Acanthocephala), Micrognathozoa, and Chaetognatha
    Phylum Gnathostomulida: The Gnathostomulids
    The Gnathostomulid Body Plan
    Phylum Rotifera: The Free-Living Rotifers
    The Rotifer Body Plan
    Body Cavity, Support, and Locomotion
    Nervous System and Sense Organs
    Reproduction and Development
    Phylum Rotifera, Subclass Acanthocephala: The Acanthocephalans
    The Acanthocephalan Body Plan
    Phylum Micrognathozoa: The Micrognathozoans
    The Micrognathozoan Body Plan
    Phylum Chaetognatha
    Chaetognath Classification
    The Chaetognath Body Plan
    Reproduction and Development
    Chapter 12
    Platytrochozoa and Two Enigmatic Phyla: Entoprocta and Cycliophora
    Phylum Entoprocta: The Entoprocts
    The Entoproct Body Plan
    Phylum Cycliophora: The Cycliophorans
    Chapter 13
    Introduction to the Lophotrochozoa, and the Phylum Mollusca
    The Lophotrochoza
    Phylum Mollusca
    Taxonomic History and Classification
    The Molluscan Body Plan
    Molluscan Evolution and Phylogeny
    Chapter 14
    Phylum Nemertea: The Ribbon Worms
    Taxonomic History and Classification
    The Nemertean Body Plan
    Nemertean Phylogeny
    Chapter 15
    Phylum Annelida. The Segmented (and Some Unsegmented) Worms
    Taxonomic History and Classification
    The Annelid Body Plan
    Sipuncula: The Peanut Worms
    Sipunculan Classification
    The Sipunculan Body Plan
    Thalassematidae: The Spoon Worms
    Siboglinidae: Vent Worms and Their Kin
    Siboglinid Taxonomic History
    The Siboglinid Body Plan
    Hirudinea: Leeches and Their Relatives
    The Hirudinean Body Plan
    Orthonectida: Extremely Simplified Annelids
    Annelid Phylogeny
    Chapter 16
    The Lophophorata: Phyla Phoronida, Bryozoa, and Brachiopoda
    Taxonomic History of the Lophophorates
    The Lophophorate Body Plan
    Phylum Phoronida: The Phoronids
    The Phoronid Body Plan
    Phylum Bryozoa: The Moss Animals
    The Bryozoan Body Plan
    Phylum Brachiopoda: The Lamp Shells
    The Brachiopod Body Plan
    Chapter 17
    Rouphozoa: The Phyla Platyhelminthes (Flatworms) and Gastrotricha (Hairy-Bellied Worms)
    Introduction to Rouphozoa
    The Phylum Platyhelminthes (Flatworms)
    Taxonomic History and Classification
    The Platyhelminth Body Plan
    Platyhelminth Phylogeny
    Phylum Gastrotricha: The Gastrotrichs, or Hairy-Bellied Worms
    The Gastrotrich Body Plan
    Reproduction and Development
    Chapter 18
    Introduction to Ecdysozoa: Scalidophora (Phyla Kinorhyncha, Priapula, Loricifera)
    Introduction to Ecdysozoa
    The Scalidophora
    Phylum Kinorhyncha: The Kinorhynchs, or Mud Dragons
    The Kinorhynch Body Plan
    Phylum Priapula: The Priapulans, or Penis Worms
    Priapulan Body Plan
    Nervous System and Sense Organs
    Phylum Loricifera: The Loriciferans
    Chapter 19
    Nematoida: Phyla Nematoda and Nematomorpha
    Phylum Nematoda: Roundworms
    Nematode Classification
    The Nematode Body Plan
    Life Cycles of Some Parasitic Nematodes
    Phylum Nematomorpha: Horsehair Worms and Their Kin
    The Nematomorph Body Plan
    Chapter 20
    Panarthropoda and the Emergence of the Arthropods. Tardigrades, Onychophorans, and the Arthropod Body Plan
    Phylum Tardigrada
    The Tardigrade Body Plan
    Phylum Onychophora
    The Onychophoran Body Plan
    An Introduction to the Phylum Arthropoda
    The Arthropod Body Plan and Arthropodization
    The Evolution of Arthropods
    Chapter 21
    Phylum Arthropoda. Subphylum Crustacea: Crabs, Shrimps, and Their Kin
    Classification of the Crustacea
    Synopses of Crustacean Taxa
    The Crustacean Body Plan
    Crustacean Phylogeny
    Chapter 22
    Phylum Arthropoda. Hexapoda: Insects and Their Kin
    The Hexapod Body Plan
    Hexapod Evolution
    Chapter 23
    Phylum Arthropoda. Subphylum Myriapoda: Centipedes, Millipedes, and Their Kin
    Myriapod Classification
    The Myriapod Body Plan
    Myriapod Phylogeny
    Chapter 24
    Phylum Arthropoda. The Chelicerata
    Synopses of Living Chelicerate Groups*
    The Euchelicerate Body Plan
    The Class Pycnogonida
    The Pycnogonid Body Plan
    Chelicerate Phylogeny
    Chapter 25
    Introduction to Deuterostomia, and the Phylum Hemichordata
    Introduction to the Deuterostomia
    Phylum Hemichordata: Acorn Worms and Pterobranchs
    The Hemichordate Body Plan
    Class Enteropneusta (Acorn Worms)
    Class Pterobranchia (Pterobranchs)
    Hemichordate Fossil Record and Phylogeny
    Chapter 26
    The Phylum Echinodermata
    The Phylum Echinodermata
    Taxonomic History and Classification
    The Echinoderm Body Plan
    Echinoderm Phylogeny
    Chapter 27
    Phylum Chordata: Cephalochordata and Urochordata
    Phylum Chordata, Subphylum Cephalochordata: The Lancelets
    The Cephalochordate Body Plan
    Phylum Chordata, Subphylum Urochordata: The Tunicates
    The Tunicate Body Plan
    Chordate Phylogeny
    Chapter 28
    Perspectives on Invertebrate Phylogeny


"“Brusca is an excellent reference text that will fill you in on material, providing you a feeling of having had a solid introduction to the clade in question. It is also now the most up to date text, which is important with the changes in taxonomy.” -Derek Rosenberger, Olivet Nazarene University "

"“Brusca is the 21st century's authoritative reference on invertebrate zoology. The most compelling feature is its comprehensiveness. It is the book that I turn to when I need to review basic biology or systematics of invertebrates.” -Steve Dudgeon, California State University Northridge "

"“[Brusca has] the highest quality figures of any texts on the subject.” -Lynne Fieber, University of Miami "

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