Ecological Studies 226 Bettina Weber Burkhard Büdel Jayne Belnap Editors Biological Soil Crusts: An Organizing Principle in Drylands Ecological Studies Analysis and Synthesis Volume 226 SeriesEditors MartynM.Caldwell Logan,Utah,USA SandraD´ıaz Cordoba,Argentina GerhardHeldmaier Marburg,Germany RobertB.Jackson Durham,NorthCarolina,USA OttoL.Lange Wu¨rzburg,Germany DelphisF.Levia Newark,Delaware,USA HaroldA.Mooney Stanford,California,USA Ernst-DetlefSchulze Jena,Germany UlrichSommer Kiel,Germany Moreinformationaboutthisseriesathttp://www.springer.com/series/86 2); 5. 2 g. Fi e e s s n o ati n a pl x e d n a d n e g e l d e ail et d or (f et n a pl ur o of nt e m p o el v e d e er h p s o bi of s ar e y of s n o milli of s d n a s u o h t g n ustsduripindler crS alsoilandF. ologic¨Budel biB. ofy nb Evolutiodrawing Bettina Weber • Burkhard Bu¨del • Jayne Belnap Editors Biological Soil Crusts: An Organizing Principle in Drylands Editors BettinaWeber BurkhardBu¨del MultiphaseChemistryDepartment PlantEcologyandSystematics MaxPlanckInstituteforChemistry DepartmentofBiology Mainz,Germany UniversityofKaiserslautern Kaiserslautern,Germany JayneBelnap SouthwestBiologicalScienceCenter U.S.GeologicalSurvey Moab,Utah USA Additionalmaterialtothisbookcanbedownloadedfromhttp://extras.springer.com. ISSN0070-8356 ISSN2196-971X (electronic) EcologicalStudies ISBN978-3-319-30212-6 ISBN978-3-319-30214-0 (eBook) DOI10.1007/978-3-319-30214-0 LibraryofCongressControlNumber:2016941282 ©SpringerInternationalPublishingSwitzerland(outsidetheUSA)2016 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilarmethodologynowknownorhereafterdeveloped. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publicationdoesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexempt fromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. 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Printedonacid-freepaper ThisSpringerimprintispublishedbySpringerNature TheregisteredcompanyisSpringerInternationalPublishingAGSwitzerland Preface Duringthelastdecades,andprobablyatleastpartlytriggeredbythefirstEcological Studies volume on this topic (volume 150), biological soil crusts (biocrusts) have quicklygainedincreasinginterestfrommanyaudiences,withwellabove100pub- licationsperyearpublishedduringthelastfewyears.Thus,wefoundthatthereis nowawealthofnewdatacoveringawiderangeofdifferenttopicsonbiocrustsand showing that biocrusts can act as “an organizing principle in drylands.” This observation inspired us to use this as the title of this second Ecological Studies volume.Thisbookisdividedintosevensectionscomprisingatotalof25different chapters. Whenweselectedtheauthorsforthedifferentchapters,wehadtwointentionsin ourmind.Onewastoincludemanyscientistsfromasmanyregionsoftheworldas possible, who also used different methodological approaches in order to get a thoroughandcomprehensiveviewonthedifferenttopics.Second,wealsowanted togetagoodmixtureofyoungerandwell-establishedresearchers.Withatotalof 61 chapter authors and 28 of them being in the doctoral, postdoc, or associate professor stage, ~46 % of the book authors are in a nonfinal/early stage of their career. Thissecondbookonbiologicalsoilcrustswouldnothavebeenpossiblewithout thehelpofsomepeopleinparticular,whomwewouldliketothankhere:firstofall, we would like to express our sincere thanks to Otto L. Lange, who gave us the opportunity to be editors of this book, who strongly supported us throughout the whole process of development, from the first ideas to the final editions, and who read and gave highly productive feedback to every single chapter of this book. Thank you so much! Second, we also would like to thank all the authors who contributedtothebook.Wearewellawareofthefactthatwesometimesaskedfor ratherprofoundandtime-consumingchangesduringthedevelopmentofthebook chapters.Despitethis,allauthorsremainedhighlycooperativeandmotivated,afact which cannot be taken for granted and which we deeply appreciate. Third, we wouldliketoexpressoursincerethankstoDr.AndreaSchlitzberger,whocoordi- natedthisbiocrustbookforSpringer.Shealwayswasextremelypatientandhelped v vi Preface usimmediatelyuponallsmallerandlargerproblems,anditreallywasapleasureto work with her. Fourth, we want to thank our supervisors and colleagues, who allowed ustospend somuchtime andenergyonthisbookandwhobelievedthat intheend,wewouldproduceaworthwhilebookthatwouldadvancethisscientific field.Finally,wealsowouldliketoexpressoursincerethankstoourfamiliesand partners for their tolerance and support during the endeavor of this second biocrustbook. We sincerely hope this book will help many scientists, land managers, policy makers, and also the environmentally interested public, to receive an overall introduction into the fascinating world of biocrusts and that it will foster many new ideas andscientific projects. Ourgoal is reached if thisbooksupports under- standingoftheoverallroleofbiocrustsasanorganizingprincipleindrylands. Mainz,Germany BettinaWeber Kaiserslautern,Germany BurkhardBu¨del Moab,UT,UnitedStates JayneBelnap Contents PartI Introduction 1 BiologicalSoilCrustsasanOrganizingPrincipleinDrylands. . . . 3 JayneBelnap,BettinaWeber,andBurkhardBu¨del 2 HowBiologicalSoilCrustsBecameRecognizedasaFunctional Unit:ASelectiveHistory. . .. . . .. . . .. . . .. . . . .. . . .. . . .. . . .. 15 OttoL.LangeandJayneBelnap PartII Morphology,Composition,andDistributionofBiological SoilCrustsatDifferentScales 3 TerrestrialEcosystemsinthePrecambrian. . . . . . . . . . . . . . . . . . 37 HugoBeraldi-CampesiandGregoryJ.Retallack 4 CyanobacteriaandAlgaeofBiologicalSoilCrusts. . . . . . . . . . . . . 55 BurkhardBu¨del,TamaraDulic´,TatyanaDarienko,NataliyaRybalka, andThomasFriedl 5 BacteriaandNon-lichenizedFungiWithinBiological SoilCrusts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 StefanieMaier,LuciaMuggia,CherylR.Kuske,andMartinGrube 6 BryophytesWithinBiologicalSoilCrusts. . . . . . . . . . . . . . . . . . . . 101 RodneyD.Seppelt,AlisonJ.Downing,KirstenK.Deane-Coe, YuanmingZhang,andJingZhang 7 Structure,Composition,andFunctionofBiocrustLichen Communities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 RogerRosentreter,DavidJ.Eldridge,MartinWestberg, LauraWilliams,andMartinGrube 8 MicrofaunaWithinBiologicalSoilCrusts. . . . . . . . . . . . . . . . . . . . 139 BrianJ.DarbyandDeborahA.Neher vii viii Contents 9 CompositionandMacrostructureofBiologicalSoilCrusts. . . . . . . 159 ClaudiaColesie,VincentJohnMartinNoahLinusFelde, andBurkhardBu¨del 10 ControlsonDistributionPatternsofBiologicalSoilCrusts atMicro-toGlobalScales. . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . 173 MatthewA.Bowker,JayneBelnap,BurkhardBu¨del, ChristopheSannier,NicolePietrasiak,DavidJ.Eldridge, andV´ıctorRivera-Aguilar 11 HypolithicCommunities. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199 StephenB.Pointing 12 RemoteSensingofBiologicalSoilCrustsatDifferentScales. . . . . 215 BettinaWeberandJoachimHill PartIII FunctionalRolesofBiologicalSoilCrusts 13 MicrostructureandWeatheringProcessesWithinBiological SoilCrusts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 237 FerranGarcia-Pichel,VincentJohnMartinNoahLinusFelde, SylvieLaureenDrahorad,andBettinaWeber 14 PatternsandControlsonNitrogenCyclingofBiological SoilCrusts. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 257 NicholeN.Barger,BettinaWeber,FerranGarcia-Pichel, EliZaady,andJayneBelnap 15 CarbonBudgetsofBiologicalSoilCrustsatMicro-,Meso-,and GlobalScales. .. . . .. . . . .. . . . .. . . . .. . . . .. . . . .. . . . .. . . . .. 287 LeopoldoG.Sancho,JayneBelnap,ClaudiaColesie,JoseRaggio, andBettinaWeber 16 BiologicalSoilCrustsasSoilStabilizers. . . . . . . . . . . . . . . . . . . . . 305 JayneBelnapandBurkhardBu¨del 17 TheRoleofBiocrustsinAridLandHydrology. . . . . . . . . . . . . .. . 321 SoniaChamizo,JayneBelnap,DavidJ.Eldridge,YolandaCant(cid:1)on, andOumarouMalamIssa 18 PhysiologyofPhotosyntheticOrganismsWithinBiologicalSoil Crusts:TheirAdaptation,Flexibility,andPlasticity. . . . . . . . . . . . 347 T.G.AllanGreenandMichaelC.F.Proctor PartIV InteractionsBetweenBiologicalSoilCrustsandVascular Plants 19 InteractionsofBiologicalSoilCrustswithVascularPlants. . . . . . . 385 YuanmingZhang,AsaL.Aradottir,MarceloSerpe, andBertrandBoeken Contents ix 20 BiologicalSoilCrustsasaModelSysteminEcology. . . . . . . . . . . 407 FernandoT.Maestre,MatthewA.Bowker,DavidJ.Eldridge, JordiCortina,RobertoLa´zaro,AntonioGallardo, ManuelDelgado-Baquerizo,MiguelBerdugo, AndreaP.Castillo-Monroy,andEnriqueValencia PartV ThreatstoBiologicalSoilCrusts 21 EffectsofLocal-ScaleDisturbanceonBiocrusts. . . . . . . . . . . . . . . 429 EliZaady,DavidJ.Eldridge,andMatthewA.Bowker 22 BiocrustsintheContextofGlobalChange. . . . . . . . . . . . . . . . . . . 451 SashaC.Reed,FernandoT.Maestre,Rau´lOchoa-Hueso, CherylR.Kuske,AnthonyDarrouzet-Nardi,MelOliver,BrianDarby, LeopoldoG.Sancho,RobertL.Sinsabaugh,andJayneBelnap PartVI NaturalandEnhancedRecoveryandManagement 23 NaturalRecoveryofBiologicalSoilCrustsAfterDisturbance. . . . 479 BettinaWeber,MattBowker,YuanmingZhang,andJayneBelnap 24 EnhancedRecoveryofBiologicalSoilCrustsAfterDisturbance. . . 499 YungeZhao,MatthewA.Bowker,YuanmingZhang,andEliZaady PartVII FutureResearchonBiologicalSoilCrusts 25 SynthesisonBiologicalSoilCrustResearch. . . . . . . . . . . . . . . . . . 527 BettinaWeber,JayneBelnap,andBurkhardBu¨del TaxonomicIndex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 535 SubjectIndex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 543