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Ecological Studies, Vol. 150 Analysis and Synthesis Edited by I.T. Baldwin, Jena, Germany M.M. Caldwell, Logan, USA G. Heldmaier, Marburg, Germany O.L. Lange, Würzburg, Germany H. A. Mooney, Stanford, USA E.-D. Schulze, Jena, Germany U. Sommer, Kiel, Germany Springer-Verlag Berlin Heidelberg GmbH Soil crust community on terra rossa soils dominated by the whitish (when dry) lichen Diploschistes diacapsis in a Mediterranean shrub community, Cyprus. Insert: The moist lichen with black apothecia (photo is close to actual lichen size). (Photo O.L. Lange) Jayne Belnap Otto L. Lange (Eds.) Biological Soil Crusts: Structure, Function, and Management 1st Edition 2001, Revised 2nd Printing 2003 With 141 Figures, 64 in Color, and 30 Tables 11 Springer Prof. Dr. Jayne Belnap US Geological Survey Forest and Rangeland Ecosystem Science Center 2290 S. Resource Blvd. Moab,Utah 84532 USA Prof. Dr. Drs. h. c. Otto L. Lange Universität Würzburg Julius-von-Sachs-Institut für Biowissenschaften Lehrstuhl für Botanik II Julius-von-Sachs-Platz 3 97082 Würzburg Germany Cover illustration: Schematic block diagram of a biological soil crust with typical colonizers (for details see Fig. 1.1 in Chap. 1). Illustration by Renate Klein-Rödder ISSN 0070-8356 ISBN 978-3-540-43757-4 Library of Congress Cataloging-in-Publication Data applied for Die Deutsche Bibliothek - CIP-Einheitsaufnahme Biological soil crusts: structure, function, and management; with 30 tables / Jayne Belnp ; Otto L. Lange (ed.). - 1. ed., rev. 2. print..- Berlin ; Heidelberg ; New York ; Barcelona ; Hong Kong ; London ; Milan ; Paris; Tokyo : Springer, 2003 (Ecological studies; Vol. 150) ISBN 978-3-540-43757-4 ISBN 978-3-642-56475-8 (eBook) DOI 10.1007/978-3-642-56475-8 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9,1965, in its current version, and permissions for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. http://www.springer.de © Springer-Verlag Berlin Heidelberg 2001, 2003 Originally published by Springer-Verlag Berlin Heidelberg New York in 2003 Softcover reprint of the hardcover 1st edition 2003 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Cover design: design & production GmbH, Heidelberg Typesetting: Friedmut Kröner, Heidelberg SPIN 10882381 31/3130 YK - 5 4 3 2 1 0 - Printed on acid free paper Preface In arid and semiarid lands throughout the world, where the cover of vegetation is sparse or absent,the open spaces between the higher plants are generally not bare ofautotrophic life,but covered by a community ofhighly specialized organisms. This soil-surface community consists of cyanobac- teria, algae, lichens, mosses, microfungi, and other bacteria in differing proportions.Cyanobacterial and microfungal filaments,rhizinae and rhizo- morphs of lichens,and the rhizinae and protonemata of bryophytes weave throughout the top few millimeters ofsoil,gluing loose soil particles together. This forms a crust up to a few centimeters thick that stabilizes and protects soil surfaces from erosive forces (Cameron and Blank 1966;Friedmann and Galun 1974;Belnap and Gardner 1993).These crusts occur in all hot,cool,and cold-arid and semiarid regions ofthe world,and may constitute up to or more than 70% of the living cover.Biological soil crusts have only recently been recognized as having a major influence on terrestrial ecosystems. Research on these soil-surface communities began in the 1950s.The first efforts were mostly descriptive, consisting of taxonomic studies of crusts from different environments.Limited research in the 1960s documented the ability of crustal components to fix nitrogen and influence hydrologic pro- cesses such as water runoffand infiltration rates.As the importance ofthese crusts in ecosystem functioning has become apparent,research efforts have intensified, with well over 3000 publications now available on the biology, ecology,and ecophysiology ofsoil-crust communities and their components, as well as on applied aspects such as landscape-level hydrology and manage- ment. Globally, this consortium of soil biota has many similarities in species composition,structure,and function,in spite of occurring in unconnected and seemingly dissimilar environments.Crusts are found in an astonishing variety of higher plant communities throughout the world: in all types of desert and semidesert plant communities, ranging from shrub deserts to succulents and to open woodlands (the latter,for instance,in the United States and in Australia). Crusts are also found in steppe formations in both the Northern and Southern Hemisphere; in the gaps of evergreen shrubs and VI Preface forests in the mediterranean-type climate;and on open ground or between the tundra vegetation in cold climates. On a small scale, soil-crust com- munities are even found in the more open types of vegetation in temperate regions,such as xerothermic local steppe formations in central Europe and in the pine barrens of the United States.Many of the dominant cyanobacteria and soil lichens have a cosmopolitan distribution, and cryptogamic com- munities ofmany crusts have almost identical growth forms.For instance,the same soil lichens dominate soil crusts of both the Sonoran and Great Basin Deserts ofthe western United States,while vascular plant species and climate varies greatly between these two regions. There are similarities in species composition of biological soil crusts even between arctic and hot desert environments.The physical structure ofcrusts is very similar in hot deserts of the world,such as the Atacama,Sonoran,Chihuahuan,African,and Australian Deserts,while very different from the physical structure ofcrusts in cool and cold deserts such as the Great Basin and the Arctic. Recently,ecophysiologists have begun to study carbon gain and nitrogen fixation ofsoil-crust biota under controlled conditions in the laboratory and in the field.These are the first steps towards understanding the soil crusts’func- tioning and their importance in different ecosystems.Researchers have found the autotrophic components of soil crusts deliver assimilates to the often carbon-limited desert soils. Cyanobacteria and cyanobacteria-containing lichens can be an important source of nitrogen for plants and soils in many desert ecosystems.In this way,crusts can improve soil fertility which,in turn, influences vascular plant nutrition.Soil crusts can also influence the germina- tion and establishment of vascular plants.Biological soil crusts reduce the susceptibility of soils to both wind and water erosion.Polysaccharides ex- truded by the cyanobacteria and green algae, along with the rhizinae of lichens and mosses,entrap and bind soil particles together,hardening the soil surface.Wind and water are major erosive forces,especially in deserts,where there is little protection of soil surfaces by organic matter or higher-plant cover.Thus,the presence ofbiological soil crusts in water-limited areas can be crucial in reducing soil loss from plant interspaces and in maintaining vascular vegetative productivity. Extensive livestock grazing, foot traffic, and off-road vehicles break through the protective soil-crust cover.This decreases the resistance of the soil surfaces to wind and water erosion.Subsequent invasion ofexotic annual plants into native perennial communities can cause dramatic changes in soil- crust flora.Such invasions,along with disturbance,can lead to substantial alterations in carbon and nitrogen inputs.Thus,land managers in arid and semiarid areas need to understand the role of crusts, and the impact of different land uses,in different ecosystems. There have been several reviews ofthe literature on biological soil crusts, beginning with Cameron and Blank in 1966 and Friedmann and Galun in Preface VII 1974.Harper and Marble reviewed the literature again in 1988,emphasizing studies in the United States, and West (1990) published a review that emphasized European and Israeli literature. This was followed by a brief overview by Johansen in 1993 and a review by Eldridge and Greene in 1994. Most recently, Evans and Johansen (1999) summarized our knowledge on biological crusts and ecosystem processes. Eldridge and Tozer (1997) published a practical guide for identification ofsoil lichens and bryophytes of Australia’s dry country. However, apparently contradictory information on the role ofthese crusts has often been published.Moreover,many papers are difficult to find,making it impossible for many people to obtain an overview on the state of knowledge about biological soil crusts.For these reasons,we felt a general synthesis volume on biological crusts was needed as a central reference for scientists and land managers.To achieve this goal,we invited the world’s leading specialists in soil-crust research to summarize known research on the different aspects ofbiological soil crusts.These contributions summarize our present knowledge about the role of soil crusts in different ecosystems,and point out known gaps in our current understanding.They also explore worldwide and regional patterns to help explain apparent differences in the role and function of these crusts from disparate environ- ments. The taxonomy of the highly specialized soil-crust organisms is a difficult field,as several ofthe systematic groups are insufficiently known for correct identification of all representatives down to the species level.Nevertheless, taxonomic composition of most of the crust types can be clearly charac- terized. In Part I of our Volume, characteristic soil-crust organisms and typical crust habitats throughout the world are depicted by photographs (Chap. 1). The communities of organisms forming soil crusts and the appearance of different soil-crust types are described for the different geographical regions.Discussion is centered on how species composition and dominance of crustal organisms varies with precipitation,temperature,soil physical and chemical properties, elevation, vascular plant vegetation, and other site factors. A synthesis chapter (Chap. 12) compares and discusses observed patterns on a continental and global scale. Biological soil crusts regularly serve as an environment also for heterotrophic soil organisms such as invertebrates and microfungi.Part IIreports what little we know about this topic.Structure and fine structure ofsoil crusts from their different habitats, using light and electron microscopy,are analyzed in the first chapter (Chap. 15) of Part III,and environmental controls on crust composition and struc- ture are discussed on various scales in the subsequent chapters. Present knowledge of crust photosynthesis and nitrogen fixation is summarized in ecophysiological chapters ofPart IV.Theinfluence ofsoil crusts on ecosystem nitrogen and carbon cycles and the effects of soil crust on vascular plant germination,survival,and nutrition is evaluated.The following parts of the VIII Preface Volume are devoted to larger-scale aspects of soil-crust performance. Re- search on the effects of soil crusts on landscape hydrology and erosion has been mainly done in Israel,Australia,and the United States.In Part V,eminent case studies are presented on soil stability and soil-water relations which include water runoff,water infiltration,evaporation and sediment produc- tion,and how these factors vary with climate,soil chemistry,soil texture and structure,as well as with crustal composition and disturbance history.Since there are seemingly contradictory findings on the role that crusts play in different soil types, geographical regions, and environments, a synthesis chapter (Chap. 26) explores regional and global patterns to explain these results.Disturbance of soil crusts can impact ecosystems in many ways,in- cluding increasing susceptibility to soil erosion and decreased carbon and nitrogen inputs. Resistance and resilience of crusts to disturbance varies widely with differences in species composition and soil types. In Part VI, impact of native and non-native ungulates,recreation,fire,and other soil- surface disturbances on species composition, biomass, and physiological functioning ofsoil crusts are reported.Ways to hasten recovery ofdisturbed areas are examined. Part VII looks at strategies to manage soil crusts, considering human impact such as off-road driving and livestock grazing. How to monitor and integrate soil crusts into ecosystem management on a land-scape scale is outlined,and use ofremote sensing for crust monitoring is represented. The possible impacts of global climate changes and how this might influence land management decisions are also discussed.Finally,Part VIII summarizes the material presented in this Volume. Resistance and resilience of soil-crust systems to disturbance are discussed, applying an evolutionary perspective.Future research needs are outlined,with manage- ment and scientific implications ofthe research summarized. As evidenced by the large and diverse audiences present at biological soil- crust workshops and seminars that have taken place in the past few years (e.g., St. Clair and Johansen 1993), land management agencies, private citizens,and the scientific community are beginning to realize the importance and exciting scientific opportunities to be found in work on biological soil crusts.As worldwide needs and pressures upon arid landscapes increase,it is also incumbent upon the scientific community to better understand the role these crusts play in different,and often fragile,arid ecosystems,and to inform land-management agencies on the impacts ofdifferent types and intensities of land use on this vital ecosystem component.We sincerely hope that the present Volume will help to procure for soil crusts the attention and apprecia- tion they deserve for the benefit of protecting arid land from long-lasting anthropogenic impacts. Due to space restrictions,only a fraction ofliterature relevant to soil-crust biology is referenced in the present Volume.One ofus (J.B.) maintains a web- based databank ofpublications concerning all aspects ofbiological soil crusts Preface IX (www.soilcrust.org).Currently (March 2001),this web site contains over 3000 references.This literature is recommended to the readers of this Volume for more detailed information regarding biological soil crusts. Acknowledgments.Dr.Burkhard Büdel (Kaiserslautern) is thanked for his continuous support and advice during the realization ofthis Volume.Soil crust research ofO.L.L. was funded under the Sonderforschungsbereich 251 der Universität Würzburg.Grateful acknowledgment is made of Beth Coker-Roy, Sue Phillips,Wilma Samfaß, Michelle Schmid,and last,but not least,Dr.Andrea Schlitzberger,who assisted in all aspects of this Volume.Friedmut Kröner provided superb production advise and expertise.Finally, we thank Rose and Steve for their patience and understanding with our absorbed state during this project. Jayne Belnap,Moab,Utah,U.S.A. March 2001 Otto L.Lange,Würzburg,Germany References Belnap J,Gardner JS (1993) Soil microstructure ofthe Colorado Plateau:the role ofthe cyanobacterium Microcoleus vaginatus.Great Basin Nat 53:40–47 Cameron RE,Blank GB (1966) Desert algae:soil crusts and diaphanous substrata as algal habitats.Jet Propulsion Lab,CalifInst Techn,Pasadena,Techn Rep 32–971:1–41 Eldridge DJ,Greene RSB (1994) Microbiotic soil crusts:a review oftheir roles in soils and ecological processes in the rangelands ofAustralia.Aust J Soil Res 32:389–415 Eldridge DJ, Tozer ME (1997) A practical guide to soil lichens and bryophytes of Australia’s dry country.Dep Land Water Cons,Sydney Evans RD,Johansen JR (1999) Microbiotic crusts and ecosystem processes.Crit Rev Plant Sci 18:183–225 Friedmann EI,Galun M (1974) Desert algae,lichens and fungi.In: Brown GW (ed) Desert biology.Academic Press,New York,pp 165–212 Harper KT,Marble JR (1988) A role for nonvascular plants in management ofarid and semiarid rangeland.In:Tueller PT (ed) Vegetation science applications for rangeland analysis and management.Kluwer Academic Publishers,Dordrecht,pp 135–169 Johansen JR (1993) Cryptogamic crusts ofsemiarid and arid lands ofNorth America.J Phycol 29:140–147 St Clair LL,Johansen JF (1993) Introduction to the symposium on soil crust com- munities.Great Basin Nat 53:1–4 West NE (1990) Structure and function of soil microphytic crusts in wildland eco- systems ofarid and semi-arid regions.Adv Ecol Res 20:179–223

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