Geobotany Studies Basics, Methods and Case Studies Elgene O. Box Kazue Fujiwara E ditors Warm-Temperate Deciduous Forests around the Northern Hemisphere Geobotany Studies Basics, Methods and Case Studies Editor Franco Pedrotti University of Camerino Via Pontoni 5 62032 Camerino Italy Editorial Board: S. Bartha, Va´cra´t(cid:1)ot, Hungary F. Bioret, University of Brest, France E. O. Box, University of Georgia, Athens, Georgia, USA ˇ A. Carni, Slovenian Academy of Sciences, Ljubljana (Slovenia) K. Fujiwara, Yokohama City University, Japan D. Gafta, “Babes-Bolyai” University Cluj-Napoca (Romania) J. Loidi, University of Bilbao, Spain L. Mucina, University of Perth, Australia S. Pignatti, Universita` degli Studi di Roma “La Sapienza”, University of Rome, Italy R. Pott, University of Hannover, Germany A. Vela´squez, Centro de Investigaci(cid:1)on en Scie´ncias Ambientales, Morelia, Mexico R. Venanzoni, University of Perugia, Italy For furthervolumes: http://www.springer.com/series/10526 About the Series Theseriesincludesoutstandingmonographsandcollectionsofpapersonagiventopicinthe followingfields:Phytogeography,Phytosociology,PlantCommunityEcology,Biocoenology, Vegetation Science, Eco-informatics, Landscape Ecology, Vegetation Mapping, Plant Con- servationBiologyandPlantDiversity.Contributionsareexpectedtoreflectthelatesttheoreti- cal and methodological developments or to present new applications at broad spatial or temporal scales that could reinforce our understanding of ecological processes acting at the phytocoenosisandlandscapelevel.Casestudiesbasedonlargedatasetsarealsoconsidered, providedthattheysupportrefinementofhabitatclassification,conservationofplantdiversity, orpredictionofvegetationchange.“GeobotanyStudies:Basics,MethodsandCaseStudies”is the successor to the journal Braun-Blanquetia, which was published by the University of Camerino between 1984 and 2011 with the cooperation of the Station Internationale de Phytosociologie (Bailleul-France) and the Dipartimento di Botanica ed Ecologia (Universita` di Camerino, Italy) and under the aegis of the Socie´te´ Amicale Francophone de Phytosociologie,theSocie´te´ Franc¸aisedePhytosociologie,theRheinoldTu¨xenGesellschaft, and the Eastern Alpine and Dinaric Society for Vegetation Ecology. This series aims to promote the expansion, evolution and application of the invaluable scientific legacy of the Braun-Blanquetschool. Elgene O. Box (cid:129) Kazue Fujiwara Editors Warm-Temperate Deciduous Forests around the Northern Hemisphere Editors ElgeneO.Box KazueFujiwara GeographyDepartment YokohamaCityUniversity UniversityofGeorgia Yokohama Athens,Georgia Japan USA ISSN2198-2562 ISSN 2198-2570(electronic) ISBN978-3-319-01260-5 ISBN978-3-319-01261-2(eBook) DOI10.1007/978-3-319-01261-2 SpringerChamHeidelbergNewYorkDordrechtLondon LibraryofCongressControlNumber:2014957871 #SpringerInternationalPublishingSwitzerland2015 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartofthematerialis concerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,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 dissimilar methodology now known or hereafter developed. Exemptedfromthislegalreservationarebriefexcerptsinconnectionwithreviewsorscholarlyanalysisormaterial suppliedspecificallyforthepurposeofbeingenteredandexecutedonacomputersystem,forexclusiveusebythe purchaserofthework.Duplicationofthispublicationorpartsthereofispermittedonlyundertheprovisionsofthe CopyrightLawofthePublisher’slocation,initscurrentversion,andpermissionforusemustalwaysbeobtainedfrom Springer.PermissionsforusemaybeobtainedthroughRightsLinkattheCopyrightClearanceCenter.Violationsare liabletoprosecutionundertherespectiveCopyrightLaw. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublicationdoesnot imply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevantprotectivelawsand regulationsandthereforefreeforgeneraluse. Whiletheadviceandinformationinthisbookarebelievedtobetrueandaccurateatthedateofpublication,neitherthe authorsnortheeditorsnorthepublishercanacceptanylegalresponsibilityforanyerrorsoromissionsthatmaybe made.Thepublishermakesnowarranty,expressorimplied,withrespecttothematerialcontainedherein. Printedonacid-freepaper SpringerispartofSpringerScience+BusinessMedia(www.springer.com) Preface Thebasicworldwidegeographicframeworkforunderstandingrelationsbetweenclimateand plant or vegetation types involves the recognition of regular bioclimatic zones, such as tropical, temperate, and further subdivisions. In global bioclimatic zonation, one zone is the warm-temperate,whichhasgenerallywarm(ifnothot)summersandrelativelymildwinters, but with some frost, which distinguishes this zone from the tropical climates. Many warm- temperateclimatesarealsogenerallyhumid,withyear-roundrainfall.Asaresult,the“zonal” (i.e., potential natural) vegetation ofthe warm-temperate zone is considered tobe evergreen broad-leaved forest, such as the “laurel forests” of East Asia, southern Brazil, and the cloud beltofsomepartsoftheCanaryIslands.Thetreestypicallyhavequitepliable,oftenglossy, thin-leatheryevergreenleaves;buildrelativelydeeprootsystems;andgrowrapidlyunderthe goodclimateconditions. In some parts of the Northern Hemisphere, though, some more “southern” deciduous temperate (summergreen) forests, mostly oak forests, extend into the warm-temperate zone, wherewintersarejustabittoocoldforthebroad-leavedevergreentrees,thoughsummersare still quite warm. That deciduous forests may occur regularly in the warm-temperate zone, undersuchconditions,wasfirstrecognizedbytheJapaneseecologistTatsuo¯ Kira,whocalled theseforests“warm-temperatedeciduous.”Thisbookrepresentsthefirstattempttorecognize and describe “warm-temperate deciduous forests” as a consistent forest type representing a consistentresponsetosimilarclimaticsituationsaroundtheentireNorthernHemisphere.Most of the chapters included herein are from papers presented in 2011, in a special session on warm-temperatedeciduousforests(alsoafirst),heldattheannualmeetingoftheInternational AssociationforVegetationScienceinLyon. Forest composition, structure, and environmental relations are treated in this book in variousways,someemphasizingbioclimaticrelationships,somefollowingtraditionalphyto- sociological analysis, plus some other methodologies. Some chapters on more specialized aspects are also included, in particular on various aspects of two prototype warm-temperate deciduous tree species, namely, downy oak (Quercus pubescens) from southern Europe and Chinese cork oak (Quercus variabilis) from East Asia. The book begins with a general overview chapter on the concept of warm-temperate deciduousness, and there is an attempt attheendtoquantifytheclimaticlimitsofpotentialtemperateforestregionsandtomapthem accordingly,aroundtheentireNorthernHemisphere. As in any large area with diverse language and cultural regions, there are problems of terminology.Thetermsusedhereinforbioclimaticzonationrepresentwhatappearstobethe global consensus on concepts, types, and names. The official Chinese national classification systemisquitedifferent,however,andsometermsareindirectconflict.Forexample,inthe Chinesesystem,alltemperatedeciduousforestshavebeencalled“warm-temperate”(despite winter temperatures to ‐30(cid:1)C or lower), and all evergreen broad-leaved forests are usually assigned to the “subtropical” zone (if not tropical). For this book, only the more globally consistent terminology is used. This is explained in the “Overview” chapter, along with an attempttoexplaintheChinesediscrepancy. Another difference involves people’s names. In East Asia the family name comes first, followed by given names. For Japanese, there is no problem, because Japanese scientists v vi Preface follow the Western order quite consistently when using Western languages. Chinese and Korean names, on the other hand, are shown much less consistently, and it has become fashionable among eager young Chinese researchers to use the Western name order consis- tently,evenwhennotdictatedbytheirpublishers.Thiscanbequiteconfusingforpeoplenot familiar with Chinese and Korean names. Family names in both Chinese and Korean are normallyofonesyllable,followedusually(butnotalways)bytwogivennames.Theproblem comeswhenthereisonlyonegivenname(onesyllable),buteventhiscanbeovercomesimply by observing the pattern used for other names in the particular publication. In this book, Chinese and Korean names are given in their natural order (family name first), with the two givennameshyphenated;JapanesenamesaregiveninWesternorder.Theonlyexceptionisin someregisterednamesofphytosociologicalsyntaxa,whichmaybeidiosyncratic. Literature references are given in the direct, common-sense format used formerly by the Ecological Society of America. In this format, the first author (of multiples) has the family namefirstandsubsequentauthorsfollowwithgivenandfamilynamesintheirnormalorder– without inversions and without all those unnecessary, obfuscating commas that make the gothic, but now “accepted” format so difficult to read. Note that, in the format used herein, there is no comma in any Chinese or Korean name because there can be no inversions of familyandgivennames. Diacritical marks are unavoidable in order to retain some correct meanings and pronunciations, especially in Japanese. For long vowels, a macron is used on o (o¯) and u (u¯),whiletheotherlongvowelsareindicated(asinwrittenJapanese)bydoublingthevowel, i.e.,aaorii,orbytheconvention“ei”forlonge.Althoughithasrecentlybecomefashionable inJapantoomitmacronsinRomanization,thedifferencebetweenlongandshortvowelsisnot trivial.Itrepresentsdifferentspellings,difference insyllablestress,andoftenquitedifferent meanings. For example, short o means small (小) but long o¯ means large (大). Other things beingequal,asyllablewithalongvowelismuchmorelikelytobestressedthanonewithonly ashortvowel–asalsoinEnglishandmostlanguages.InChinese,eachsyllablecarriesoneof four possible tones. Tone marks, though, are usually not shown herein, since these do not representdifferencesinspelling,eitherinpinyin(theChinesephoneticRomanizationsystem) orinWesternrenditions. Finally, understanding some place names, especially in China, will be much easier if the reader is willing to learn just three East Asian words. A mountain (or mountains) is shan in ChineseandsaninbothJapaneseandKorean;ling(Chinese)isarangeofmountains.Thus, [the]Changbai-ShanisthemountainsalongtheborderbetweenChinaandNorthKorea,Fuji- sanisMt.Fuji,andtheDaXingAnLing(Chinese)istheGreaterHingganmountainrangethat separatesInnerMongolia(tothewest)fromChineseManchuria(totheeast). SadlywemustalsonotethatTatsuo¯ Kirapassedawayin2011,atage91.Heoncetoldme (Box)thathekepthisimaginationyoungbydividinghisscientificcareerintoroughly10-year segments,ineachoneofwhichhestudiedsomethingentirelydifferentfrombefore.Perhaps this is a good model for many of us – his life was a model for all of us. It seems fitting, therefore,thatwededicatethisbooktohismemory. Athens(Georgia),USA ElgeneO.Box Yokohama,Japan KazueFujiwara Contents Introduction:WhyWarm-TemperateDeciduousForests?. . . . . . . . . . . . . . . . . . 1 ElgeneO.BoxandKazueFujiwara Warm-TemperateDeciduousForests:ConceptandGlobal Overview. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 ElgeneO.BoxandKazueFujiwara CharacterofWarm-TemperateQuercusForestsinAsia. . . . . . . . . . . . . . . . . . . . 27 KazueFujiwaraandAtsukoHarada PhytosociologicalStudyofQuercusvariabilisForestin Warm-TemperateChina. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81 TangQian,KazueFujiwara,andYouHai-Mei Warm-TemperateForestsofCentralPortugal:AMosaic ofSyntaxa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 Jose´ CarlosCosta,TiagoMonteiro-Henriques,PedroBingre, andDalilaEsp´ırito-Santo MarcescentForestsoftheIberianPeninsula:Floristicand ClimaticCharacterization. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119 ItziarGarc´ıa-Mijangos,JuanAntonioCampos,IdoiaBiurrun, MercedesHerrera,andJavierLoidi Downy-OakWoodsofItaly:PhytogeographicalRemarksona ControversialTaxonomicandEcologicIssue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 RiccardoGuarino,GiuseppeBazan,andBrunoPaura TheStatusofQuercuspubescensWilld.inEurope. . . . . . . . . . . . . . . . . . . . . . . . 153 CamillaWellsteinandFrancescoSpada OnthePotentialofQuercuspubescensWilld.andOtherSpeciesof QuercusintheCamerinoSyncline(CentralItaly). . . . . . . . . . . . . . . . . . . . . . . . . 165 KevinCianfaglione ChorologyandPhytosociologyofSessileOak[Quercuspetraea (Mattuschka)Liebl.]inTrentino-AltoAdige(East-CentralAlps) ofNorthernItaly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175 FrancoPedrotti QuercussuberDistributionRevisited. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181 BartolomeoSchirone,FrancescoSpada,MarcoCosimoSimeone, andFedericoVessella PhenorhythmsandForestRefugia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213 BartolomeoSchirone,FrancescoSpada,GianlucaPiovesan, andMarcoCosimoSimeone vii viii Contents Warm-TemperateDeciduousForestsofEasternNorthAmerica. . . . . . . . . . . . . . 225 ElgeneO.Box DeciduousWoodlandsinMediterraneanCalifornia. . . . . . . . . . . . . . . . . . . . . . . 257 MarkA.Blumler DeciduousWoodlandsintheNearEasternFertileCrescent, andaComparisonwithCalifornia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267 MarkA.BlumlerandJoelC.Plummer QuantitativeDelimitationofWarm-TemperateDeciduous ForestAreas. . .. . . .. . . . .. . . .. . . . .. . . .. . . . .. . . .. . . .. . . . .. . . .. . . . .. . 277 ElgeneO.Box Appendix:ClimaticDataforRepresentativeHumid-TemperateSites AroundtheNorthernHemisphere,withClimateTypePredictedfrom anEnvelopeModelandMapped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285 Introduction: Why Warm-Temperate Deciduous Forests? Elgene O. Box and Kazue Fujiwara Warm-temperate deciduous forests would seem to be an Inthissense,warm-temperatedeciduousforestscanbecon- enigma. In the various global (and globally aware) bio- ceivedas: climatic zonation systems, warm-temperate climates are – “southern”temperatedeciduousforests; generally construed as humid temperate climates with – occurring where summers are warm but winters are too warm summers, mild winters (no lasting snow cover), and coldforevergreens; evergreen broad-leaved forest as the zonal (i.e. climatic – occurringnotonlyinJapanbutalsomainlandEastAsia, potential) vegetation type. This is the general conceptual- southeasternUSA,andperhapselsewhere;and ization both in global treatments of vegetation and climate, – including perhaps the “thermophilous” and asbyRu¨bel(1930),Schmithu¨sen(1968),andWalter(1968, “submediterranean” deciduous forests of southern 1970, 1985); and in regional systems, such as by Kira and Europe. othersforEastAsia(cfKira1945,1949,1977,1991;Suzuki Why do such deciduous forests persist in warm-temperate 1953; Miyawaki 1967). Such zonal evergreen broad-leaved areas?Aretheyanomalies,oraretheyanaturalalternativein forests occur in East Asia (often called “laurel forests”) as aconsistentglobalzonation? well as in southern Brazil, northern New Zealand, parts of Warm-temperate deciduous forests (and more open easternAustralia,inmontanebeltsoftropicalAsia(andeven woodlands) are a Northern Hemisphere phenomenon, theCanaryIslands),and,wheretopographypermits,insmall resulting, like the boreal forest, from the continentality of areasofthewarm-temperatesoutheasternUSA.Thermally, the large Northern land masses (cf Troll 1948; Box 2002). some mediterranean climates are also warm-temperate and Kira recognized the warm-temperate deciduous forests of may carry evergreen broad-leaved forests, albeit with Japan by observing that some deciduous Quercus species sclerophyllratherthanlaurophyllfoliage. inJapandonotoccurineitheroftheadjacentclimaticzones, Even so, Tatsuo¯ Kira (1949) recognized that deciduous thecool-temperatetothenorthorthesubtropicalzonetothe forests may also occur as stable forests in some warm- south. Indeed,throughout the NorthernHemisphere, warm- temperate areas, if winters are too cold for evergreens. temperatedeciduousforestsareoverwhelminglyoakforests. For these forests, especially some in interior Honshu¯ (the In East Asia these forests are composed especially of deci- largestislandofJapan),hecoinedtheterm‘warm-temperate duous Quercus serrata, Q. variabilis, Q. aliena and Q. deciduous forest’. These deciduous forests occur not only dentata. These species form forests in the drier climates of as successional stages but may remain as permanent vege- interiorHonshu¯,suchasaroundNagano,wheresummersare tation, even in some areas where winters are milder. warm but winters are colder. Minimum winter temperature in these areas can go below (cid:1)15(cid:3)C (not below (cid:1)20(cid:3)C), mean January temperature is around 0(cid:3)C, and values of Kira’s Coldness Index are below (cid:1)10(cid:3)C. Average annual precipitationmaybe900–1100mm,whichislowforJapan. Similar deciduous forests also occur in the relatively E.O.Box(*) warm climates of north-central to eastern China and the GeographyDepartment,UniversityofGeorgia,Athens,Georgia30602, more southern lowlands of Korea; analogous deciduous USA forests beyond Asia appear to include the Quercus e-mail:[email protected] pubescens forests of the Mediterranean area and the oak- K.Fujiwara hickory (Quercus-Carya) forests of interior southeastern YokohamaCityUniversity,Yokohama236-0027,Japan e-mail:[email protected] North America. In East Asia, the main warm-temperate E.O.BoxandK.Fujiwara(eds.),Warm-TemperateDeciduousForestsaroundtheNorthernHemisphere, 1 GeobotanyStudies,DOI10.1007/978-3-319-01261-2_1,#SpringerInternationalPublishingSwitzerland2015