Ecology. Michael Begon

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      ECOLOGY

      From Individuals to Ecosystems

       MICHAEL BEGON

       University of LiverpoolLiverpool, UK

       COLIN R. TOWNSEND

       University of OtagoDunedin, New Zealand

      FIFTH EDITION

      This fifth edition first published 2021 © 2021 by John Wiley & Sons Ltd

      Edition History 4e, Wiley, 2006; 3e, Wiley, 1996; 2e, Wiley 1990; 1e, Wiley, 1986

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      The right of Michael Begon and Colin R. Townsend to be identified as the authors of this work has been asserted in accordance with law.

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       Library of Congress Cataloging‐in‐Publication Data

      Names: Begon, Michael, author. | Townsend, Colin R., author.

      Title: Ecology : from individuals to ecosystems / Michael Begon, The University of Liverpool, Liverpool, UK, Colin Townsend, University of Otago, Dunedin, New Zealand.

      Description: Fifth edition. | Hoboken, NJ : Wiley, 2021. | Includes bibliographical references and index.

      Identifiers: LCCN 2020012775 (print) | LCCN 2020012776 (ebook) | ISBN 9781119279358 (paperback) | ISBN 9781119279372 (adobe pdf) | ISBN 9781119279310 (epub)

      Subjects: LCSH: Ecology.

      Classification: LCC QH541 .B415 2020 (print) | LCC QH541 (ebook) | DDC 577–dc23

      LC record available at https://lccn.loc.gov/2020012775 LC ebook record available at https://lccn.loc.gov/2020012776

      Cover Design: Wiley

      Cover Credit: Decorators painting wall © Dougal Waters/Getty Images

      Panel of Native American Newspaper Rock petroglyphs © milehightraveler/Getty Image

       This book is dedicated to our families, and especially to our children and grandchildren,who will inherit the ecosystems we hand on to them –by Mike to Linda, Jessica, Robert, Carl, Ria and Erica,and by Colin to Dominic, Brennan, Amelie and Ella

      Preface

      A science for everybody – but not an easy science

      This book is about the distribution and abundance of living organisms, and about the physical, chemical and especially the biological features and interactions that determine these distributions and abundances.

      Unlike some other sciences, the subject matter of ecology is apparent to everybody: most people have observed and pondered the natural world. In this sense most people are ecologists of sorts. But ecology is not an easy science. It must deal explicitly with three separate levels of the biological hierarchy – the individual organisms, the populations of organisms, and the communities of populations. What’s more, as we shall see, it can’t ignore the detailed biology of individuals, or the pervading influences of historical, evolutionary and geological events. It feeds on advances in our knowledge of biochemistry, behaviour, climatology, plate tectonics and so on, but it feeds back to our understanding of vast areas of biology too. One of the fathers of modern evolutionary biology, T. H. Dobzhansky famously claimed, in the middle of the 20th century, that ‘Nothing in biology makes sense, except in the light of evolution’. But equally, very little in evolution, and hence in biology as a whole, makes sense, except in the light of ecology.

      Ecology, too, has the distinction of being peculiarly confronted with uniqueness: millions of different species, countless billions of genetically distinct individuals, all living and interacting in a varied and ever‐changing world. Imagine doing chemistry if water molecules were not precisely the same, always and everywhere – or physics if the speed of light was different at different field sites. The challenge for ecologists, therefore, is to seek patterns and predictions in a way that recognises this uniqueness and complexity, rather than being swamped by it. The mathematician and philosopher Alfred North Whitehead’s advice for scientists, offered 100 years ago, is never more apposite than when applied to ecology: ‘Seek simplicity – and distrust it’.

Thirty‐four years on: the urgent problems facing us

      This fifth edition comes fully 14 years after its immediate predecessor and 34 years after the first edition. Much has changed over that time – in ecology, in the world around us, and also, unsurprisingly, in us authors. The first edition had a cave painting as its cover, which we justified in the Preface by arguing that