Invasive Plant Ecology in Natural and Agricultural Systems, 2nd Edition This page intentionally left blank Invasive Plant Ecology in Natural and Agricultural Systems, 2nd Edition Barbara D. Booth Independent Consultant, Guelph, Ontario, Canada Stephen D. Murphy Department of Environment and Resource Studies, University of Waterloo, Waterloo, Ontario, Canada and Clarence J. Swanton Department of Plant Agriculture, University of Guelph, Guelph, Ontario, Canada CABI is a trading name of CAB International CABI Head Office CABI North American Office Nosworthy Way 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail: [email protected] E-mail: [email protected] Website: www.cabi.org © CAB International 2010. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Booth, Barbara D. (Barbara Diane), 1961- Invasive plant ecology in natural and agricultural systems / Barbara D. Booth, Stephen D. Murphy, and Clarence J. Swanton. -- 2nd ed. p. cm. -- (Modular texts) First edition published by CABI in 2003 with title: Weed ecology in natural and agricultural systems. Includes bibliographical references and index. ISBN 978-1-84593-605-1 (alk. paper) 1. Invasive plants--Ecology. 2. Weeds--Ecology. I. Murphy, Stephen D. II. Swanton, Clarence J. III. C.A.B. International. IV. Title. V. Series: Modular texts. SB611.B59 2011 632'.5--dc22 2010009986 First edition published by CAB International in 2003. ISBN-13: 978 0 85199 528 1 ISBN-13: 978 1 84593 605 1 Commissioning editor: Rachel Cutts Production editor: Fiona Chippendale Typeset by SPi, Pondicherry, India. Printed and bound in the UK by Cambridge University Press, Cambridge. Contents Preface vi 1 Introduction to Invasion Ecology 1 2 The Distribution and Abundance of Populations 8 3 The Structure and Dynamics of Populations 25 4 Sexual Reproduction 46 5 Asexual Reproduction 58 6 From Seed to Seedling 71 7 Growing Up, Getting Old and Dying 85 8 Competition 97 9 Herbivory, Parasitism and Mutualism 110 10 Basic Community Concepts and Diversity 124 11 Community Dynamics: Succession and Assembly 141 12 Landscape Scales and Invasive Species 164 13 Molecular Ecology: Applications for Invasive Plants 177 14 Plant Invasions: a Synthesis 186 Glossary 198 Subject Index 203 Species Index 211 v Preface to the 2nd Edition Our goal in writing this book was to describe how and why plant invasions occur. The book attempts to explain the ecological principles that are important in understanding the potential for a species to become invasive. We think students should understand the ecology of invasive plants and how this knowledge can be used to facilitate management. Ecology is central to our understanding of how and why plant species invade, and yet there are few books that make this connection. This is the niche we hope to fill. The book was designed as a teaching text for advanced undergraduate students. No extensive ecological background is assumed, although some basic biology is required. At the beginning of each chapter, we have listed the concepts that will be addressed. These can be used as an overview of what is to come, and to assist the reader when reviewing the material. For the instructor, we have designed this book so that the material could be covered in a single-term course by covering approximately one chapter per week. We have tried to write the book and arrange the material so that it is presented in a clear, precise and concise manner, and to include only pertinent information. If we have done our job well, students should be able to read and under- stand all the information. We have used common names throughout the text, with Latin names given the first time that the species is mentioned in each chapter. We did this because common names are easier to remem- ber when first learning about a species. A species list of common and Latin names is provided at the end of the book. At the end of each chapter there will be questions that refer to an invasive plant species of your (the stu- dent’s) choice (i.e. a case study). The plant can be an established invasive species, or perhaps a new species that may have potential to invade your local community You will be asked to summarize information that is known about your plant species in relation to the material discussed in each chapter. There may be a lot or very little information available to you. The idea behind this is to apply the ecological principles you learn in the chapter to an invasive species of interest, and to give you practice researching a topic. Our hope is that, by the end of the book, you will have created a case history of your chosen plant. We have made a number of changes to this second edition from the first, which was published by CABI in 2003 as Weed Ecology in Natural and Agricultural Systems. We have removed the chapters that focused solely on experimental methods and added chapters on the newer fields of landscape and molecular ecology. In addition we have updated the literature. We thank our publisher CABI for their help and encouragement with this second edition. Of course we accept the responsibility for any errors that occur. Finally, we thank our spouses, David Beattie, Tara Murphy and Josee Lapierre, who once again throughout the writing of this second edition heard way more about ‘the book’ than they ever wanted, but kept smiling and nodding their heads anyway. We dedicate this second edition to them. vi Introduction to Invasion Ecology 1 Producing and EatCionncgep tMs eat ● The terms invasive, colonizer, weed and others are often used in overlapping and conflicting manners. ● An invasion is the geographical expansion of a species into an area not previously occupied. ● Both native and non-native species can be invasive. ● Most invasions fail. ● The impact of an invasion depends on the area covered by the species, its abundance and the effect per individual. ● Invasion meltdown is the acceleration of impacts on native ecosystems due to synergistic interactions. 1.1 Introduction invasive species (Sheley and Petroff, 1999; Myers and Bazely, 2003; Liebman et al., 2007; Clout and It may be tempting for you to start this book Williams, 2009), or provide a detailed description with Chapter 2. After all, the real information of the biology of individual invasive species with- doesn’t start until then, and exam questions out providing a broad background (Czarapata, rarely focus on what you learn in Chapter 1. 2005; Young and Clements, 2009). Here we focus BUT Chapter 1 is important because it sets the on the underlying ecological principles that explain tone for what is to follow. A Shakespearean play the process of plantinvasions. or an opera always begins with a prologue. If There are a number of excellent plant ecology you walk in after the prologue has finished, you texts (e.g. Barbour et al., 1999; Gurevitch et al., will certainly follow the plot and enjoy the play, 2006). This book was not designed to replace a but you might not understand the ‘why’ of the good, comprehensive text on basic ecological the- characters’ actions. Consider this chapter a pro- ory. Rather, its purpose is to examine invasive plant logue. You may already know much of what we species in systems from agricultural and natural are about to say, and you may not be tested on communities. Whether an invasive plant is in a it, but it will put what you are about to learn natural community or a highly managed farm, the into context. underlying questions and principles will be the In this book, our goal is to provide you with a same. All types of ecological systems are controlled basic ecological understanding of how plants by the same processes. invade natural, disturbed and agricultural ecosys- tems. Whether an invasive plant is in a natural community or a highly managed farm, the under- 1.2 Levels of Ecology lying questions and principles will be the same. A number of books are available on plant inva- The word ecology was derived from the German sions; however, they often assume an in-depth word (oekologie), which was derived from the understanding of ecological principles (Mooney Greek words oikos, meaning ‘house’, and logos, et al., 2005; Sax et al., 2005; Cadotte et al., 2006), meaning ‘the study of’. Thus, ecology is the study of focus heavily on the control and management of organisms and their environment. We can divide the ©CAB International 2010. Invasive Plant Ecology in Natural and Agricultural Systems, 1 2nd Edition (B. Booth et al.) environment into biotic (living) and abiotic (non- ● an alien in a semi-natural habitat (Stirton, 1979; living) factors. Examples of biotic factors are com- Macdonaldet al., 1989) petition and herbivory. Abiotic factors can be ● a native or non-native entering any new habitat physical (e.g. temperature, light quality and quan- (Mack, 1985; Gouyon, 1990) tity), or chemical (e.g. soil nutrient status). ● a native or non-native that is increasing in popu- The field of ecology is vast. It is concerned with lation size (Joenje, 1987; Mooney and Drake, areas as diverse as the dispersal of seeds, competi- 1989; Le Floch et al., 1990) tion within and between species, and nutrient ● any non-native increasing in population size cycling through ecosystems. Each of these oper- (Prach and Wade, 1992; Binggeli, 1994; ates on a different temporal and spatial scale and Rejmánek, 1995) each has a different focus. Thus, they all ask dif- ● any non-native species (Kowarik, 1995). ferent types of questions, and require a different Colautti and MacIsaac (2004) summarized more protocol to answer them. For convenience, eco- recent definitions of an invasive species as: logical questions can be categorized into different scales: ● a non-native species (Goodwin et al., 1999; ● Molecular ecology examines how DNA is Radford and Cousens, 2000) ● a native or non-native species that has colonized expressed as traits that may allow plants to natural habitats (Burke and Grime, 1996) invade. ● Individual organisms can be studied to examine ● a widespread non-native species (van Clef and Stiles, 2001) how abiotic factors affect their physiology and ● a widespread non-native species that has a neg- phenology. ● Groups of individuals of the same species can be ative effect on habitat (Davis and Thompson, 2000; Mack et al., 2000). studied to look at population-level processes. ● Groups of co-occurring populations of different Clearly, there is little consensus on what invasive species can be studied to ask community-level means. Similar problems occur with other terms, questions. such as weed. Weeds have typically been defined as ● Interactions between a community and its abiotic ‘plants which are a nuisance’ (Harper, 1960) or ‘a factors can be studied to answer landscape- and plant where we do not want it’ (Salisbury, 1961). ecosystem-level questions. Barbour et al. (1999) defined a weed as a ‘non- native invasive plant’ and they distinguished between Each of these categories blends into the next. ‘invasive plants’ that invade only natural or slightly They are not discrete units of study, rather they disturbed habitats, and ‘pest plants’ that interfere are useful, practical and somewhat arbitrary with agricultural or managed natural areas. This divisions which help to simplify the understand- definition, however, requires us to further define ing of invasion ecology. In this book we discuss non-native and invasive, and to separate natural population ecology (Chapters 2–7), interactions from disturbed habitats. These definitions are based among populations (Chapters 8 and 9), commu- on our perceptions of the impact of the plant. nity ecology (Chapters 10 and 11), landscape Similarly, the terms invader and colonizer have often ecology (Chapter 12), and molecular ecology been used in a conflicting manner. The distinctions (Chapter 13). In the final chapter (14) we incor- between them are quite subtle and result from differ- porate all levels of ecology to discuss the process ing viewpoints. According to Rejmánek (1995), of plant invasions. weeds interfere with human land use; colonizers are successful at establishing following disturbance; and invaders are species introduced into their non-native 1.3 What’s in a Name? habitat. There is substantial overlap among these Every book on invasive species must first start terms. A plant may be considered as only one of with an attempt at defining terms. Many attempts these, or it may be included in all of these categories have been made to define invasive and other terms (Fig. 1.1). Given the confusion of terminology, we describing a species’ status. Pyšek (1995), for take a broad view of the term invasive, defining it as example, reviewed definitions of an invasive spe- a species that has a negative ecological or economic cies and found that it has been described as: effect on a natural or managed ecosystem. 2 Chapter 1 Anthropogenic viewpoint: Ecological viewpoint: interferes with objectives or pioneers in succession requirements of people Weeds Colonizers e z si e g Invaders n a R Biogeographical viewpoint: introduced, exotic, alien, non-native species Fig. 1.1. Weeds, colonizers, and invaders are similar concepts but result from differing viewpoints (redrawn from Rejmánek, 1995). Time It is important to differentiate species that are Fig. 1.2. The process of invasion. invasive from species that are introduced. While most invasive species are introduced, this is not size will level off. In Chapters 2 to 13 of this text always the case. For example, in the north-eastern we will discuss individual processes that influence USA, the native hay-scented fern (Dennstaedtia one or several of these stages of invasion, and in punctilobula) becomes invasive under conditions of the final chapter we synthesize these individual intensive browsing by white-tailed deer and the chapters and revisit the invasion process. thinning or removal of overstorey tree canopy (de la Cretaz and Kelty, 1999). In addition, not all introduced species become invasive. Many plant Most invasions fail, but some will succeed species are intentionally introduced for use as Most invasions fail. That is, most species when crops, for erosion control, or for aesthetic reasons introduced into a community will not survive. (e.g. in gardens). While some introduced species A successful invasion is a rare event (Williamson, have become invasive, many will persist in small 1996). How rare? Williamson (1996) proposed self-sustaining populations but will not expand in the ‘tens rule’ to describe how approximately range or will remain at low density. Furthermore, 10% of species pass through each transition from many introduced species would go extinct in their being imported (dispersal) to becoming casual introduced range without human management (e.g. (introduced to the wild) to becoming established, maize will never become invasive in Canada). and finally becoming an invasive (Williamson 1996) (Fig. 1.3). This is a rough rule, but has been 1.4 What is an Invasion? shown to apply around the world, under many situations (Williamson and Fitter, 1996). What is Elton (1958) was the first person to formalize the evident from this rule is that successful invasions study of invasion ecology, recognizing that inva- are rare (Williamson, 1996). sions of introduced species could have global Our knowledge of why an invasion fails can be impacts (Richardson and Pyšek, 2008). Since quite limited (Rejmánek, 1999). Successful invaders then, interest in invasion ecology has grown. An are obvious, but failed invaders are not; therefore, invasion is ‘the geographical expansion of a spe- it is difficult to study the process of a failed inva- cies into an area not previously occupied by that sion. Some reasons that invasions fail are (Crawley, species’ (Vermeij, 1996). The process that a plant 1987; Lodge, 1993): species goes through when it invades has a series of stages. Initially, following a species’ introduc- ● inappropriate abiotic conditions for the species tion, its geographical expansion occurs very to survive slowly (Fig. 1.2). This lag period can last as long ● introduced species are outcompeted by other as centuries. Following the lag period, there species may be exponential expansion of the species’ ● presence of natural enemies such as herbivores range. Eventually, the increase in a species’ range and diseases Introduction to Invasion Ecology 3