Monica G. Turner Robert H. Gardner Landscape Ecology in Theory and Practice Pattern and Process Second Edition L ANDSCAPE E COLOGY IN T HEORY AND P RACTICE M ONICA G . T URNER R OBERT H . G ARDNER LANDSCAPE ECOLOGY IN THEORY AND PRACTICE Pattern and Process Second Edition Monica G. Turner University of Wisconsin-Madison Department of Zoology Madison , WI , USA Robert H. Gardner University of Maryland Center for Environmental Science Frostburg, MD , USA ISBN 978-1-4939-2793-7 ISBN 978-1-4939-2794-4 (eBook) DOI 10.1007/978-1-4939-2794-4 Library of Congress Control Number: 2015945952 Springer New York Heidelberg Dordrecht London © Springer-Verlag New York 2015 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifi cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfi lms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specifi c statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. Printed on acid-free paper Springer Science+Business Media LLC New York is part of Springer Science+Business Media (www.springer.com) Dedication Frank B. Golley: Scholar, Mentor, Friend here are people whom we encounter in life who make a real difference. TThey may infl uence us as individuals by their encouragement and guidance; they may affect their community through their untiring commitment to service and the greater good; or they may infl uence an entire profession by their scholarship, leadership, and example. We dedicate this second edition of Landscape Ecology in Theory and Practice to Frank B. Golley (1930–2006) who made a real difference in all of these ways. Frank spent most of his career at the University of Georgia and was Monica’s graduate advisor. We marked his many accomplishments previously (see Turner et al. 2007, Landscape Ecology 22:1–3) but note here his service as founding editor of Landscape Ecology , president of the Ecological Society of America and the International Association for Ecology (INTECOL), and director vi of the Biological Sciences Directorate at the National Science Foundation. But beyond his many academic achievements and the lasting infl uence he had on the development of landscape ecology, Frank was a wonderful person, a mentor, and friend who touched both of us deeply. We would not be where we are today with- out him. We thank him for all that he gave of himself, and we still mourn his loss. Frank, this is for you. Preface to the Second Edition Heterogeneity, pattern, process, scale, and hierarchy are intrinsically interrelated concepts … key to the theory and practice of landscape ecology. Wu (2013) he literature of landscape ecology has expanded tremendously since the fi rst edition of Landscape Ecology in Theory and Practice was fi nished in T early 2000. Indeed, the number and diversity of published landscape studies is stag- gering. It is wonderful that the fi eld has blossomed, but synthesizing the advance- ments and literature of landscape ecology has been a very daunting task. Nonetheless, we have tried to capture key developments in the fi eld, but we also recognize that this goal will probably be met only partially. Despite a large bibliog- raphy with well over 1000 references cited, there was no way to include all studies. By necessity, our examples and citations are illustrative, not comprehensive. How is the second edition organized, and in what ways does it differ from the fi rst edition? The fi rst two chapters set the stage. Chapter 1 ( Introduction to Landscape Ecology and Scale ) introduces the fi eld of landscape ecology and reviews its intellectual foundations, and it now includes a basic introduction to scale con- cepts and terminology. Scale remains important, but we no longer felt it required a stand-alone chapter; however, scale issues infuse the rest of the book. Chapter 2 ( Causes of Landscape Pattern ) has been much enhanced. Treatment of multiple and contingent causation is much expanded, along with the challenges associated with forecasting future landscape patterns in the context of multiple changing drivers. viii Landscape development and landscape legacies are also treated in greater depth, refl ecting advances in understanding of how land-use history affects contemporary landscapes. After setting the stage, three chapters deal with quantitative methods in land- scape ecology. These analysis methods are key tools for scholars and practitioners in landscape ecology and the means to the end in many studies. Chapter 3 ( Introduction to Models ) focuses on simulation modeling, providing an overview of spatially explicit landscape models, which are used widely in landscape ecology. Chapter 3 also presents neutral landscape models (NLMs), which were instrumen- tal in early quantitative studies of pattern–process relationships. Some readers (especially those who are new to spatial pattern analysis) may prefer to read Chaps. 4 and 5 before reading Chap. 3 . Chapter 4 ( Landscape Metrics ) covers the range of landscape metrics used with categorical data, including metrics calculated for the entire landscape and by cover type or patch. This chapter has been updated to introduce some newer approaches, but as in the past, it emphasizes proper use and interpretation of metrics rather than exhaustive coverage. Chapter 5 ( Spatial Statistics ) is new and was needed because of substantial advances in methods for analyzing spatial dependence in continuous data, and their wide use. With the foundation established and quantitative methods presented, the next three chapters deal with particular pattern–process relationships. All three chapters have been substantially revised and updated to refl ect the growth of knowledge in these areas. Chapter 6 ( Disturbance Dynamics ) now includes new topics such as landscape epidemiology and disturbance interactions. Chapter 7 ( Organisms and Landscape Patterns ) was extended to cover interactions among species, community structure, species invasions, and landscape genetics. Each of these topics has seen tremendous progress in the past 15 years. Chapter 8 , Ecosystem Processes in Heterogeneous Landscapes ) distinguishes between ecosystem processes character- ized primarily by vertical fl uxes and processes that represent lateral transfers. The last two chapters consider applications and synthesis. Chapter 9 ( Landscape Dynamics in a Rapidly Changing World ) deals with applications of landscape ecol- ogy and has been thoroughly revised. This chapter now considers landscape indica- tors, climate change, and landscape sustainability, in addition to an expanded treatment of land-use change and landscape scenarios. The fi nal and relatively brief chapter ( Conclusions and Future Directions ) summarizes key lessons from land- scape ecology and directions for future research. Acknowledgements Research that we have conducted over the past 15 years has contributed to the writing of this book. Turner especially acknowledges research funding in support of this effort from the National Science Foundation, including the Long-Term Ecological Research Program (Coweeta LTER: DEB-0823293 and North Temperate ix Lakes LTER: DEB-0822700) and Water Sustainability and Climate Program (DEB- 1038759); the Joint Fire Science Program (Grants 09-1-06-3, 09-3-01-47, 11-1-1- 7); the Andrew W. Mellon Foundation; the Department of Zoology and Graduate School of the University of Wisconsin-Madison; and the Wisconsin Department of Natural Resources. Turner thanks all the students and postdocs who have been part of my lab for many spirited and fun discussions about landscape ecology; I learn so much from you. I especially thank my recent lab members Pete Blank, Dan Donato, Catherine Frock, Rose Graves, Winslow Hansen, Brian Harvey, Jiangxiao Qiu, Kevin Rose, Amy Uhrin, Tim Whitby, and Carly Ziter, not only for their input but also their understanding when I have absented myself to write. My ideas and understanding of landscape ecology have also benefi tted from long-term collaborations with val- ued colleagues, especially Bill Romme and Scott Pearson, and the outstanding group of excellent ecologists at the University of Wisconsin-Madison. Graduate students in my Advanced Landscape Ecology course at UW-Madison have been a tremendous source of inspiration and new knowledge. I also thank Sarah Gergel for her hard work and perseverance to complete the second edition of Learning Landscape Ecology , which makes a great companion to this text and offers many options for hands-on learning of the approaches we describe here. Gardner thanks the faculty, staff, and students of the Appalachian Laboratory for their generous support and enthusiasm during the development of this new edi- tion. Special thanks are due Katia Engelhardt, Andrew Elmore, and Matt Fitzpatrick for stimulating discussions, collaboration, and assistance in teaching landscape ecology. None of this would be possible without the love and support of wife, Kim S. Gardner, and family. The University of Maryland Center for Environmental Science (UMCES) is a unique institution of which the Appalachian Laboratory is a part. Faculty and students of other UMCES labs, especially the Horn Point Laboratory and Chesapeake Biological Laboratory, provided a stimulating envi- ronment for vetting concepts and ideas with particular thanks to Michael Kemp and Walter Boynton who listened critically and freely offered new ideas.