PHENOLOGY: AN INTEGRATIVE ENVIRONMENTAL SCIENCE

Tasks for vegetation science 39

SERIES EDITORS

A. Kratochwil, University of Osnabrück, Germany H. Lieth, University of Osnabrück, Germany

The titles published in this series are listed at the end of this volume.

PHENOLOGY: An IntegrativeEnvironmental Science

Edited by

MARK D. SCHWARTZDepartment of Geography,University of Wisconsin – Milwaukee, Milwaukee, WI, U.S.A.

KLUWER ACADEMIC PUBLISHERS

DORDRECHT / BOSTON / LONDON

A C.I.P. Catalogue record for this book is available from the Library of Congress.

ISBN 1-4020-1580-1

Published by Kluwer Academic Publishers,P.O. Box 17, 3300 AA Dordrecht, The Netherlands.

Sold and distributed in North, Central and South Americaby Kluwer Academic Publishers,101 Philip Drive, Norwell, MA 02061, U.S.A.

In all other countries, sold and distributedby Kluwer Academic Publishers,P.O. Box 322, 3300 AH Dordrecht, The Netherlands.

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All Rights Reserved© 2003 Kluwer Academic PublishersNo part of this work may be reproduced, stored in a retrieval system, or transmittedin any form or by any means, electronic, mechanical, photocopying, microfilming, recordingor otherwise, without written permission from the Publisher, with the exceptionof any material supplied specifically for the purpose of being enteredand executed on a computer system, for exclusive use by the purchaser of the work.

Printed in the Netherlands

Dedication

This book is dedicated to myparents, Marjorie H. and thelate Donald J. Schwartz, who nurtured my early interest in

science

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Contents

Contributing Authors xi

Preface xvii

Color Plates xxi

Foreword xxvii

Part 1: INTRODUCTION 1

1.1 Introduction 3MARK D. SCHWARTZ

Part 2: PHENOLOGICAL DATA, NETWORKS, AND RESEARCH 9

2.1 East Asia 11XIAOQIU CHEN

2.2 Australia 27MARIE R. KEATLEY AND TIM D. FLETCHER

Dedication v

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2.3 Europe 45ANNETTE MENZEL

2.4 North America 57MARK D. SCHWARTZ AND ELISABETH G. BEAUBIEN

2.5 South America 75L. PATRÍCIA C. MORELLATO

2.6 The Global Phenological Monitoring Concept 93EKKO BRUNS, FRANK-M. CHMIELEWSKI, AND ARNOLD J. H. VANVLIET

2.7 Toward a Multifunctional European Phenology Network 105ARNOLD J. H. VANVLIET AND RUDOLFRR S. DEGROOT

Part 3: PHENOLOGY OF SELECTED BIOCLIMATIC ZONES 119

3.1 Tropical Dry Climates 121ARTURO SANCHEZ-AZOFEIFA, MARGARET E. KALACSKA, MAURICIO

QUESADA, KATHRYN E. STONER, JORGE A. LOBO, AND PABLOARROYO-MORA

3.2 Mediterranean Climates 139DONATELLA SPANO, RICHARD L. SNYDER, AND CARLA CESARACCIO

3.3 Grasslands of the North American Great Plains 157GEOFFREY M. HENEBRY

3.4 High Latitude Climates 175FRANS E. WIELGOLASKI AND DAVID W. INOUYE

3.5 High Altitude Climates 195DAVID W. INOUYE AND FRANS E. WIELGOLASKI

Part 4: PHENOLOGICAL MODELS AND TECHNIQUES 215

4.1 Plant Development Models 217ISABELLE CHUINE, KOEN KRAMER, AND HEIKKI HÄNNINEN

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4.2 Animal Life Cycle Models 237JACQUES RÉGNIÈRE AND JESSE A. LOGAN

4.3 Phenological Variation of Forest Trees 255ROBERT BRÜGGER, MATTHIAS DOBBERTIN, AND NORBERT KRÄUCHI

4.4 Phenological Growth Stages 269UWE MEIER

4.5 Assessing Phenology at the Biome Level 285XIAOQIU CHEN

4.6 Developing Comparative Phenological Calendars 301REIN AHAS AND ANTO AASA

4.7 Plant Phenological "Fingerprints" 319ANNETTE MENZEL

4.8 Phenoclimatic Measures 331MARK D. SCHWARTZ

4.9 Weather Station Siting 345RICHARD L. SNYDER, DONATELLA SPANO, AND PIERPAOLO DUCE

Part 5: REMOTE SENSING PHENOLOGY 363

5.1 Remote Sensing Phenology 365BRADLEY C. REED, MICHAEL WHITE, AND JESSLYN F. BROWN

Part 6: PHENOLOGY OF SELECTED LIFEFORMS 383

6.1 Aquatic Plants and Animals 385WULF GREVE

6.2 Insects 405KAREN DELAHAUT

6.3 Birds 421TIM H. SPARKS, HUMPHREY Q. P. CRICK, PETER O. DUNN, ANDLEONID V. SOKOLOV

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6.4 Timing of Reproduction in Large Mammals 437ERIC POST

Part 7: APPLICATIONS OF PHENOLOGY 451

7.1 Vegetation Phenology in Global Change Studies 453nMICHAEL A. WHITE, NATHANIEL BRUNSELL, AND MARK D.SCHWARTZ

7.2 Phenology of Vegetation Photosynthesis 467LIANHONG GU, WILFRED M. POST, DENNIS BALDOCCHI, T. ANDYBLACK, SHASHI B. VERMA, TIMO VESALA, AND STEVE C. WOFSY

7.3 Radiation Measurements 487JIE SONG

7.4 Phenology and Agriculture 505FRANK-M. CHMIELEWSKI

7.5 Winegrape Phenology 523GREGORY V. JONES

7.6 Long-Term Urban-Rural Comparisons 541CLAUDIO DEFILA AND BERNARD CLOT

Acknowledgments 555

Index 557

Contributing Authors

Aasa, Anto, Institute of Geography, University of Tartu, Tartu, Estonia

Ahas, Rein, Institute of Geography, University of Tartu, Tartu, Estonia

Arroyo-Mora, Pablo, Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT, USA

Baldocchi, Dennis, Department of Environmental Science, Policy & Management, University of California, Berkeley, CA, USA

Beaubien, Elisabeth G., Devonian Botanic Garden, University of Alberta, Edmonton, Alberta, Canada

Black, T. Andy, Faculty of Agricultural Sciences, University of BritishColumbia, Vancouver, Canada

Brown, Jesslyn F., SAIC, USGS EROS Data Center, Sioux Falls, SD, USA

Brügger, Robert, PHENOTOP, Institute of Geography of the University of fBerne, Berne, Switzerland

Brunns, Ekko, Department of Networks and Data, German Meteorological Service, Offenbach, Germany

Brunsell, Nathaniel, Department of Civil Engineering, Duke University,Research Triangle, NC, USA

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Cesaraccio, Carla, Agroecosystem Monitoring Laboratory, Institute of Biometeorology, National Research Council, Sassari, Italy

Chen, Xiaoqiu, Department of Geography, College of Environmental Sciences, Peking University, Beijing, China

Chmielewski, Frank-M., Subdivision of Agricultural Meteorology, Instituteof Crop Sciences, Faculty of Agriculture and Horticulture, Humboldt-University, Berlin, Germany

Chuine, Isabelle, CEFE-CNRS, Montpellier, France

Clot, Bernard, Biometeorology, MeteoSwiss, Zürich and Payerne, Switzerland

Crick, Humphrey Q. P., British Trust for Ornithology, Thetford, UK

Defila, Claudio, Biometeorology, MeteoSwiss, Zürich and Payerne,Switzerland

deGroot, Rudolf S., Environmental Systems Analysis Group, Department of Environmental Sciences, Wageningen University, Wageningen, The Netherlands

Delahaut, Karen, Department of Horticulture, University of Wisconsin- Madison, Madison, WI, USA

Dobbertin, Matthias, WSL Swiss Federal Institute for Forest, Snow andLandscape Research, Forest Ecosystems and Ecological Risks Division, Birmensdorf, Switzerland

Duce, Pierpaolo, Agroecosystem Monitoring Laboratory, Institute of Biometeorology, National Research Council, Sassari, Italy

Dunn, Peter O., Department of Biological Sciences, University of Wisconsin-Milwaukee, Milwaukee, WI, USA

Fletcher, Tim D., Department of Civil Engineering, Monash University, Clayton, Victoria, Australia

Greve, Wulf, German Center for Marine Biodiversity Research(Senckenberg Research Institute), Hamburg, Germany

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Gu, Lianhong, Environmental Sciences Division, Oak Ridge National Laboratory, Oak Ridge, TN, USA

Hänninen, Heikki, Department of Ecology and Systematics, University of Helsinki, Helsinki, Finland

Henebry, Geoffrey M., Center for Advanced Land Management InformationTechnologies (CALMIT), School of Natural Resources, University of Nebraska, Lincoln, NE, USA

Inouye, David W., Department of Biology, University of Maryland, College Park, MD, USA

Jones, Gregory V., Department of Geography, Southern Oregon University,Ashland, OR, USA

Kalacska, Margaret E., Earth and Atmospheric Sciences Department, University of Alberta, Edmonton, Alberta, Canada

Keatley, Marie R., School of Resource Management, University of Melbourne, Creswick, Victoria, Australia

Kramer, Koen, Alterra, Department of Ecology and Environment,Wageningen University, Wageningen, The Netherlands

Kräuchi, Norbert, WSL Swiss Federal Institute for Forest, Snow andLandscape Research, Forest Ecosystems and Ecological Risks Division,Birmensdorf, Switzerland

Lobo, Jorge A., Biology Department, Universidad de Costa Rica, San Jose,Costa Rica

Logan, Jesse A., USDA Forest Service, Logan, Utah, USA

Meier, Uwe, Federal Biological Research Center for Agriculture andForestry, Braunschweig, Germany

Menzel, Annette, Department of Ecology, TU Munich, Freising, Germany

Morellato, L. Patrícia C., Departmento de Botânica, Plant Phenology andSeed Dispersal Research Group, Universidade Estadual Paulista, São Paulo, Brasil

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Post, Eric, Department of Biology, The Pennsylvania State University, University Park, PA, USA

Post, Wilfred M., Environmental Sciences Division, Oak Ridge NationalLaboratory, Oak Ridge, TN, USA

Quesada, Mauricio, Centro de Investigaciones en Ecosistemas, Universidad Nacional Autónoma de México, Morelia, México

Reed, Bradley C., SAIC, USGS EROS Data Center, Sioux Falls, SD, USA

Régnière, Jacques, Natural Resources Canada, Canadian Forest Service, Quebec, Canada

Sanchez-Azofeifa, Arturo, Earth and Atmospheric Sciences Department, University of Alberta, Edmonton, Alberta, Canada

Schwartz, Mark D., Department of Geography, University of Wisconsin-Milwaukee, Milwaukee, WI, USA

Snyder, Richard L., Department of Land, Air, and Water Resources,University of California, Davis, CA, USA

Sokolov, Leonid V., Russian Academy of Sciences, St. Petersburg, Russia

Song, Jie, Department of Geography, Northern Illinois University, Dekalb, IL, USA

Spano, Donatella, Department of Economics and Woody Plant Ecosystems,University of Sassari, Sassari, Italy

Sparks, Tim H., Centre for Ecology and Hydrology, Monks Wood, UK

Stoner, Kathryn E., Centro de Investigaciones en Ecosistemas, UniversidadNacional Autónoma de México, Morelia, México

vanVliet, Arnold J. H., Environmental Systems Analysis Group, Department of Environmental Sciences, Wageningen University, Wageningen, TheNetherlands

Verma, Shashi B., School of Natural Resource Sciences, University of Nebraska, Lincoln NE, USA

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Vesala, Timo, Department of Physical Sciences, University of Helsinki,Helsinki, Finland

White, Michael, Department of Aquatic, Watershed, and Earth Resources, Utah State University, Logan, UT, USA

Wielgolaski, Frans E., Department of Biology, University of Oslo, Oslo,Norway

Wofsy, Steve C., Department of Earth and Planetary Sciences, Harvard University, Cambridge, MA, USA

Preface

I recall as a doctoral student at the University of Kansas discussing dissertation topics with my advisor, Prof. Glen A. Marotz, one day in 1983. He had just suggested to me that phenology was an interesting topic, and onethat held promise for important research contributions in the future. “What’sthat?” I asked, thus beginning my career as a phenologist, and the long path that led to my editorship of this volume.

Skipping ahead a decade, I was encouraged by my colleague Elisabeth Beaubien to attend the 13th International Congress of Biometeorology, which was being held in Calgary, Alberta, Canada that year. I did attend, and alsomet Prof. Dr. Helmut Lieth there for the first time. I had corresponded with him while writing my dissertation, having gained much insight from his seminal book, Phenology and Seasonality Modeling at that time. At thegCalgary meetings Prof. Lieth helped Elisabeth and I reactivate a PhenologyStudy Group within the International Society of Biometeorology (ISB).

The first workshop of the new group was organized by Dipl.-Met. Hartmut Scharrer of the German Weather Service (DWD) Phenology unit,and scheduled for May 1995. As a UW-Milwaukee assistant professor in the Geography Department at the time, I had never traveled outside of North America, and further did not have a source of travel funding for the trip to Offenbach (just outside Frankfurt), Germany. So I consulted the associatedean responsible for our department, G. Richard Meadows (now Dean of theCollege of Letters and Science) and he was able to provide me with funds to cover the airfare (after I assured him that this trip would be an important one for establishing my connection to international phenological research). At the Offenbach workshop the thirteen participants proposed an organizational structure and laid out a set of objectives for the Phenology Study Group.

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Some subsequent early activities included the launching of a new journal Phenology and Seasonality (unfortunately discontinued after one issue), and participation in the 14th International Congress of Biometeorology(Ljubljana, Slovenia) in 1996. The study group’s first international scientificmeeting was a “Phenology Symposium” that I organized in 1998 as a groupof four paper, one poster and one discussion sessions (21 participants) heldwithin the Association of American Geographers Annual meeting in Boston, MA, USA.

The number of individuals involved with, amount of research being conducted in, and level of interest by scientists from other disciplines for phenology had all been slowly rising since the early 1990s. However, a series of papers published in Nature (over the 1997-2000 period) dramatically accelerated these trends, especially the interest of global changeresearchers in remote sensing and biology for phenological data and techniques. In recent years, this surge in interest from the global change research community, and corresponding funding by the European Union of several projects (POSITIVE and EPN, European Phenology Network) have led to a greater number of scientific conferences with increasing numbers of participants. Specifically, a first “stand alone” international phenology conference, organized by Dr. Annette Menzel and colleagues (2000, Freising, Germany, 70 participants), and two subsequent international conferences (organized by Arnold vanVliet and associates) held inWageningen, The Netherlands in 2001 and 2003 connected with the EPN project (each had just over 100 participants). The two European projectshave also supported a large number of workshops on specialized phenology topics for smaller groups of participants (including individuals from other parts of the world). Within the ISB, the study group participated in the 15th

International Congress of Biometeorology (held in Sydney, Australia in 1999) and due to reorganization within the society was renamed theVegetation Dynamics, Climate, and Biodiversity Commission after that meeting. Members of the new ISB commission also participated in the most recent ISB Congress (16th International Congress of Biometeorology, held in Kansas City, MO, USA in 2002). During that meeting the group requested, and was subsequently granted by ISB, the simplified current name“Phenology Commission.”

So the sequence of events I have described created the conditions andprovided resources to make development of this book possible, namely sufficient interest in the topic by the general scientific community, and an interconnected community of phenological researchers with the necessary diversity of research expertise to cover the range of required topics. JaccoFlipsen, a Kluwer editor, who wrote me a letter in early 2001 stating theneed for and asking if I was interested in editing a book on plant phenology,

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initiated the actual development of this volume. After some negotiation,specifically to allow the book to cover a broader range of phenological topics, the project began in earnest during the first months of 2002. The book was seamlessly transferred into Prof. Lieth’s “Tasks for Vegetation Science” series at Kluwer (supervised by Helen Buitenkamp) in early 2003, and completed later that year.

Mark D. Schwartz

Milwaukee, March 2003

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Foreword

I was pleased when Mark Schwartz invited me to write a foreword tohis volume. And even more so after I had read the content and many of the papers contributed to the volume. My own book on the subject matter (Lieth1974) appeared as vol. 8 in the famous ecological studies series by Springer, and a contribution to the then fully operating Analysis of Ecosystemprogram of the U.S. International Biological Program (US-IBP).

Phenology was a rather quiet scientific objective at that time. Some operational networks existed in Europe and America mainly in agriculture. Only a few researchers in biology, ecology and meteorology were using theaccumulated datasets at that point. Satellite image analyses and the development of new remote sensing techniques were of interest then, but theground truth observation of biological fluctuating phenomena were regarded as outmoded.

The common thrust of the papers presented at the 1972 phenologysymposium of the American Institute of Biological Sciences conference in Minneapolis gave phenology work in the U.S. and Europe a big push, and ground truth observations in ecosystems studies were initiated in many partsof the world. The initial successes in modeling phenological events, thecomparisons with meteorological parameters, and the correlation attempts with global remote sensing data sets caught the attention of the scientists, working at that time on global change initiated directly and indirectly byhumans.

This interest continues, and a book presenting the achievements of the last 30 years (two or three generations of graduate students) is very muchneeded. I followed the results with interest, because I had earlier madepredictions that had to be tested, verified or modified through field

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observations. Phenological observations and experiments undertaken during the last 30 years have greatly improved insights into ecosystems operation. One of the major values for phenological data is their validation value for seasonality models. These models have gained prominence in global climatic change models to predict biosphere responses to climatic parameter changes.

This, however, is by no means the only value of phenological work. The book presented here by Dr. Schwartz includes many other fields of biology for which phenological investigations are needed. The reliance of species association in ecosystems upon a quasi-correct seasonal behavior in a seasonal climate is so prominent, that most investigations and experiments include phenological aspects, be they climatic, physiologic or biochemical.

Throughout the historical development of phenology, its practical applications in agriculture and forestry have dominated the field. Thechapters in this volume dealing with the history of phenology by Menzel(Chapter 2.3) and Chen (Chapter 2.1) uncovered many local networks that I had not found in the early 1970s. While this is a valuable addition to the field, I found that several important networks and papers had still been neglected. The 1974 volume has, therefore, not completely lost its relevancefor future generations of phenologists. The history of European phenologyemphasizes agricultural and forest phenology and neglects the body of work started by Heinrich Walter, whose students and coworkers (e.g., Kreeb and Ellenberg) and these together with their coworkers made substantialcontributions to phenology in Europe (see Walter 1960, which shows that he had much more influence on phenology than providing the widely used climate diagrams which are so easily available in the climate diagram worldatlas by Walter and Lieth 1960ff., and now available on CD by Lieth et al. 1999).

The Russian work on phenology is only partly recognized. For me a major omission appears to be the book by Alexander Podolski, which appeared about 2 decades ago in English (1984). His approach toidentifying the start of a phenologically valid period from physiological data, rather than an arbitrary chosen, convenient calendar date, still warrants further analyses in relevant cases. Podolski’s volume also includes a wealth of literature otherwise not mentioned in Russian books that mostly refer to papers from west Russian institutes (covered by Dr. Menzel’s historical treatment in this volume).

It appears to me that students interested in phenology should beencouraged to read some of the older papers by Hopkins (1938), Thornthwaite (several papers), Hopp, Caprio, Schnelle and Volkert, and all athe others as cited in Lieth (1974) and in this volume, as well as Walter (1960) and Podolski (1984). The literature on remote sensing and globalchange applications is so new, that for the purpose of this book’s users, the

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authors in this area of research will be available in current relevant journals. Many authors of these papers will not include their contribution as part of phenology, but their work deals very often with topics that would be included in seasonality, climate and species fluctuations, global change and methods for the investigation of these topics. All this is phenology in the wider sense. The historical assessment in another 30 years will evaluate the importance of these authors and developments for phenological work. Work on satellite remote sensing had just started around the time I compiled my phenology book. The same was true for computer mapping, which was in itsinfancy as well. But the combination of data available from different phenological networks in the U.S. through computer modeling and computer mapping was so attractive to graduate students, that many of them choose phenological topics for their degree papers.

When I developed my volume in the early 1970s I was greatlysupported by Forrest Stearns who was a professor at the University of Wisconsin-Milwaukee. Wisconsin was an intellectual center for phenology at that time where the Lettaus (Heinz and Katharina) provided guidance inmeteorological and phenological observations. No wonder then that phenology received new impulses from Wisconsin. In summary I can saythat this book edited by Dr. Schwartz shows that phenology is as alive andimportant as ever. Like any other field of research it undergoes peaks and valleys in recognition. As long as planet earth tumbles around the sun, therewill be ecologists and meteorologists, foresters and agronomists, insurancepeople and a wealth of other specialists observing, measuring and evaluating phenological data. Many of them will use this book.

I thank also the responsible persons in Kluwer academic publishers for their interest in presenting this volume with the usual Kluwer quality. I am sure that the book will obtain the worldwide reception accorded many of other previous volumes of the T:VS series.

Helmut Lieth

Osnabrück, February 2003

REFERENCES CITED

Hopkins, A. D., Bioclimatics–A science of life and climate relations, U.S. Dept. Agr. Misc.Publ. 280, 1938.

Lieth, H., editor, Phenology and Seasonality Modeling, Springer-Verlag, New York, 444 pp., 1974.

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Lieth, H., J. Berlekamp, S. Fuest, and S. Riediger, Climate Diagram World Atlas on CD(unpaginated electronic publication), Backhuys Publishers, Leiden, Netherlands, 1999.

Podolski, A. S., New Phenology: Elements of mathematical forecasting in ecology, John Wiley and Sons, New York, 504 pp., 1984.

Walter, H., Grundlagen der Pflanzenverbreitung, part 1 Standortslehre, Eugen Ulmer Verlag, Stuttgart, Germany, 566pp., 1960.

Walter, H. and H. Lieth, Klimadiagramm-Weltatlas (unpaginated), VEB Gustav Fischer, Jena, 1960ff.