Studies on the Limestone Flora of Japan and Taiwan

r「 STUDIES ON THE LIMESTONE FLORA OF JAPAN AND TAIWAN PART I Tatemi SHIMIZU BiolOgical Laboγatory， the Faculty（ゾTθ雇だθS（ゼ伽Cθand T召chnology， Shinshn Univerε露゜砂，こleda， JaPan （Received September 10，1962） CONTENTS Introduction．＿，＿．．……・…………・…・……………・… …・・………・…・∴………‘……・・1 Chapter I History of the studies……一一・・……………一・…一…………・・…3 §1 Floristic and vegetational researches ………・……………齢’・”°’”闇’’”e’°3 §20th…el・t・d p・・bl・m・…………一…・………・・………・……一……’12 Ch、pt。・II Di・t・ibuti。・・f th・1im・・t…f1・1d・i・J・p・n・nd T・iw・・…’°””17’ Ch、pt。・III Lim・・t・n・1・・al f1・ra・・fJ・p・n・・d T・iw・n…一……’…’…20． Hokkaido……・一・一………一…・…・…・…一一・・一・………・・…・・……20 H・耳・h・……………・…………一…・……一…・一・…一………”…23 』＄蜘k・……・……・一………………・一一・一…・一…・…・……°’°”…70． Ky。・hu－…・……・…………一…・・………一一一・・……・…………「’…80 Taiwan＿＿＿…・………・………………………・…・一…一……・………92 Literatures for Patt工・・…………・・………・…・…・…・…・・……………・……・・………・°98 、 INTRODUCTION ！ So far as the limestρne flora is concerned， it js well known that there exist more or less characteristic species to be called ‘‘calcicolous”or ‘‘calci－ phi1。…．The・t・di…nlimest・n・．且・ras・r v・g・t・ti・n・ca・b・・a・tly f・u・d i。many且・・i・ti…ec・1・gical literature・especi・1iy． @f・。m E・・9pe s｝nce the middle of the last century． In Japall the works dealing with thls．subject hlave been swelled in number especially quite recently． My study orl the limestone且ora was commenced in 1954． Up to the 餌esent， m。・t lim・・t…di・t・i・t・i・J・p・n were vi・it・d・F・・m N・v・mb・・1960 to May 1961，0n the other hand， I stayed in Taiwan and could make some botanical trips to the Iimestone districts there． Through these travels， there have been carried out the floristic researches together with the field observa－ ti・n・， by whi・h the ch・・adt・・i・ti…fth・1imest・ne且・・a c・・ld be rec・9・i・rd・ At the same time，皿y study was sometimes accompanied by the field work m the sense of phytosociology． It aims a vegetational comparison between limestone fields and the others， by which detection of peculiarity of the limestone communities will be possible． For analysis of speciation， cytological proce． 2 Tatemi SH【MIzu No．30 dures will present a usefu1皿ethod as weU as examination of the morpholQgical variations of plants． Whether limestone plays a r61e for chromosome aber． ration or not may be also qn interesting problem， though I have gained Iittle results yet． Basing upon these directions of approach， th呈s text is designed to be consisted of the following matters， dealing with the limestone且ora in Japan and Taiwan・as the whole and with its biological significance in the various senses as mentioned above． For a status of publication， it will be divided into three parts． PART I and PART II are concerned with the且oristic problems ill main． Chapter I and「Chapter II are the introductory part for them， mentioning research history（Chapter I）and distribution of limestone fields in Japan and Taiwan（Chapter II）． Each Ioca1且ora is summarized in Chapter III． PART I inclu．des these three chapters， Chapter IV deals with grouping of the且or呈stic components of the limestone districts and its corres． ponding exemplification． From the taxonomical and phytogeographical view－ point， the note－worthy components are discussed one by one in Chapter V， The gelleral characteristics of the limestone flora in Japan and Taiwan together with comparison with serpentine且oras are mentioned in Chapter VI． PART II includes these three chapters， PART III is prepared for the phytogeographicai qnd phytosociological proble皿s and for consideration of the general biological aspects referring to variation and speciatiQn of plants in the limestone且elds． The word“1imstone且ora”used in this text is an inclusive term meaning the flora on the habitats derived from calcareous rocks， viz． calcite， chalk， dolo面te， gypsum， limestone， marble， etc． So far as my own works are concer1ユed， however， oniy the fields of limestone or crystalline limestone will be matter of discussion． The present work has been carr三ed out mainly at the University of Kyoto under the guidance of Prof． S． KITAMuRA and Dr， M， TAGAwA． So I must mention my cordial thanks to them at the first place． It should be written specially that the studies of the serpentine floras of Japan done by Prof， S， KITA・ MuRA provided the present study with numerous suggestive accounts． In these two years， my study has been promoted thanks to kilnd direction anCl encou・ rage皿ent of Dr． N． KoYAMA of Shinshu University訊 During my stay in Taiwan， I was皿uch indebted to Prof． T． S． Lエu and Prof． C， E． DEVoL of the National Taiwan University for their P’rovision with’facilities of the study， So my special thanks must be due to theln． At the same time I wish to．express my hearty thanks to many other botanists of Japan and Taiwan who helped me ill various ways． The materials on which the present study is based are ill Inajor量ty pre・ served in the herbarium of the University of Kyoto， and part1y of Shinshu University and in others． No．30 Studies on the lirnestone fliora of’ Japan and Taiwan 3 CHAPTER I HISTORY OF THE STUDIES The plants fro卑1imestone丘elds provide the botanists with many interesting ．problems． Above all， owing to its peculiarities， researches of the limestone flora have attracted many botanists of the world． Indeed．㌻hey have the very inception al士eady as long． ago as 1836， when UNGER pointed out that the lime・．stone flora was of much peculiarity in the northern Tirol of Austria， alld grouped its mernbers into three classe呂according to their preferences for limestone． LouDoN（1838）and THuRMANN（1849）made some interestipg ol⊃servations on the relatidn between soil condition and plant distributign・ According tq．the former， a beech tree， Fagus sylvatica， which is usually growing on dry soils as chalk and limestone， is absent from west England even in calcareolユs rock ．range because of higher rainfall there and lower porosity of limestone than lchalk． The latter in turn was interested in the fact that the above beech was usually calcicolous in the llortherrしEurope while rather silicicolous irl the Meditteranean side． This phenomenon seems to have led THuRMANN to an opinion that the so－called calcicolous plants are only xerophilQus， for driness く）fthe calcareous soils is though七to pro丘t the xerophilous plants under the climate of higher humidity in north Europe than that in the Meditterane母n side． These observations will give some important suggestions for considera－ tion of the nature of the calcicolous species． Thus the investigation of limestone且oras originated o1ユone hand in．the static aspect lgoking for its Horistic peculiarities or characteristic species and ・on． the other l， hand in the dyna皿ic aspect dealing with the distributiQnal mechanism of：the calcicolous plants． III this chapter， I intend．室o summarize ．the progress ill rese，arch in these two directions’of approache忌 for convini－ ence． Needless to mention， however， the characteristics of any fiora and mechanisエnS controlling them should be pursued hand in hand． §1， Floristic．又and vegetation乱l researches Study of the limestone flora， gen． erally speakiug， seems to ha▽e been carried out in proportion to the degree of progress in the general floristic research． In Europe and North America， numerous botanists have anllounced the limestone floras as what they were， while in、the other parts of the world， apart from Japan， there are few works dealing with the subject．． EUROPE：It may be English botanists who best contributed to the limestone 丑ora． They have made many informations on the旦oras or vegetations of the ．famQus chalk land in particular， which forms an elusive horizon in south・ eastem England． Following to the genera玉explanation of HuDsQN（1900）or TANsLEY et a1（1911）， on olle hand， aseries of the clqse studies of the chalk ．vegetati6n was published by ADAMsoN （1922）， TANsLEY （1922）， TANsLEY and ADAMsoN（1925，1926）， ANDERsoN（1927）and HopE－SIMpsoN（1940，1941 a b）， in ．whi¢h they clarified the composition of． chalk vegetati叩with regard to 丑owering plants， bryophytes and lichens， o血tlin60f the plant suqcesion on 4 Tatemi SHIMIzu No．30・ chalk lands， and chemical or physical na加re of chalk downs， i． e． water content， reaction， and so on． WATT（1923，1924，1925∫1934 a b， and 1926）， ou the other hand，． studied the beech woods（Fagus syんa彦ica）and the yew woods （7「axusろaccata）on chalk lands mainly．from the view point of plant succession， SALIsBuRY（1918）， ANDERsoN（1928）， HopE－SIMpsoN（1938）， LocKET（1946）aud others may be the other investigators of chalk vegetatiorls around lthere， while BR酬cHLEY（1912）， FRAsER（1939）and HARvEY（1939）are specialists on the chalk flora rather in the western part of England including Wilts， Somerset and Devon． Depending upon these outcomes several popular books were alsol issued with references to chalk Iands；for example， TANsl，EY（1925）， GILMouR・（1947），LousELY（1950）and SALIsBuRY（1952 b）． Sumniarizing all of these works， the chalk lands in England may be outlined as follows． Edaphologically the、 soils derived from chalk， which is extremely porous rock composed of 95～99 ％CaCO3， are shallow， quickly drying and with a strongly alkaline reaction． The soil type is very si皿ilar over the elltire chalk． outcrops alld imparts a corresponding uniformity to the vegetatio1ユ． Floristically， the chalk lands are． characterized by ： Aceras anthrii）o」クhoru〃z， Aブuga cha〃lae．Pitys，！lnacamPtis’ P夕7a〃zida”S，ノlnthy〃S vulneraria，！1sPerula cynaf・zchica， Baアtsia odoフztites， Bra∬icaα1わα， B7∫za〃iedia， Ast7agalus danicas，． Brachツカodium力勿％α如鋭， B70mUS・ erecttes，．Zi｝uxus 8θ〃ゆθ7z／irensi， Camψam‘la glomerata2， Cα〃ina vulgaris， Centauriar〃1：襯ゐθ〃α’蹴，CirsiZtm aea〃θ， C．副0伽rum var．δ吻nnica， Dα％C％s Cαrota・， EUPh7asiaαタzglica，

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