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(1)253. The Present SituatiOn of Speleo10gy ln. Japanl)'2). Masuz0 1UЁ N03). (With One Plate and Ten Text‐ lgures) 1.. A Brief HistOry. Thirty_eight years ago, when l was a pOst_graduate in zo010gy at KyOto lmperial University, I was wOrking on freshwater bi010gy, and. in particular on the taxOnomy and ec010gy Of Cladocera and other freshwater C)rustacea. Naturally, I came to be interested in subterranean. species and this led me tO go to caves to coHect themo. l began with. an exp10ration of the aquatic fauna Of Akiyoshi_do (or the Akiyoshi. CaVe)in 1927, and was successful in cOHecting three species of Crustacea, One of which was registered as a new species of the isOpod genus cα θθグ グθ′ θ α (it belongs tO the genus」. tsθ ′ ′ %s at present). It Was probably the flrst scientiflc survey of cave aquatic fauna in Japan,. though twO years before Fo SILVESTRI inVestigated the terrestrial fauna 1)This is an enlarged and revised form of my lecture originally delivered before the meeting of the National Speleological Society, under the presidency of Mr. Russell H. GuRNEE, WhiCh was held On September 4, 1962, at Closter, NeⅥ″Jersey (U. So A。 ). Paragraph 4 and a bibliOgraphy at the end of this. article have been newly prepared. 2) For the preparation Of the present inanuscript, I am much indebted to Dr. Shun― Ichi uttNO, :Departinent of Z001ogy, National Science Museurrl, T6ky6, fOr. his erective assistance in furnishing me with much useful material.. My. cordial thanks are also due to Mr. Tadashi KuRAMOTO, Of the Akiyoshi― dai Science Museum (Ⅳ Ir.IsaO KAWAsAKI,Director),whO Was kind enough to arord me every facility,when l visited Akiyoshi‐ d6 in January 1965, and to Yoshiyasu. KANETsuNA WhO kindly supplied me with his unpublished data On diatoms of Akiyoshi‐ dO.. I am alsO grateful tO Mro IIirosi YAMAuTI fOr providing me with. the flgure of ]Ry色 6-dO.. 3)K6nan WOmen's College;fOrmerly Ky6to University。.

(2) Speleology in Japan. 254. of the same caveo. For my caVe studies, I gained a great deal of. inforrrlation from A.S.PACKARD'S rた (publiShed in 1888)。. 力 z422zθ ″グ εα θ cα υι Fα %π α げ Ⅳ θγ′. At that tilne no other scientiflc cave investiga‐. tions had been made in Japan; only a few weH_known caves, such as Akiyoshi― dO, were visited by tourists 、rith curiosity. In 1935, when my text‐ bcok. On freshwater biology was published in the JapaneSe language,. I reviewed the progress of researches on subterranean fauna, in vaters by referring to the work on anilnal life of subterranean 、. Hermann SPANDL and Pierre A. CHAPPUIS (bOth b00kS similarly θπ Gθ zυ assθ ″),aS Well as to Rene ″ググ sε 力 ″グθ″ %%′ ιγグ entitled」 Dグ θ Tグ θγzυ θノ α F″ α″εθ. After that tilne, however, εθ′ ι グθ ノ JEANNEL'S Fα %″ 形 θαυι″πグ. nO speleological activities had become. ■ourishing in Japano My interest. in research work then turned to limnological investigations of lakes,. including both plankton and benthic anilnals, as weH as physical and cherrlical properties of lake waterS.. After the War, our knowledge of the fauna of JapaneSe caves increased rapidly, thanks largely to the works by J. ISHIKAWA, H. TORII, R.YosII, Shun‐ Ichi UЁ NO and Some others, particularly to con‐ siderable research by Professor YosII and the last‐. named.. Besides. these biospeological studies, we owe much to the fundamental work. by Mr. Hirosi YAMAUTI, Assiatant Professor at Ehilne University, Matsuyama in Shikoku.. Having overcome many initial difnculties in. exploration, he has. brought a great many caves to light and has. prepared a large number of rnaps of these caves. A number of Japanese. biologists and other scientists who were interested in speleological ished to organize a society for the purpose of promot― work,ardently 、ア. ing research activities on caves and subterranean waters.. Then, in. 1954, the Spelaeological Society of Japan was founded and l was appointed its presidente. At present, there are in Japan four speleo_. logical organizations, of which two are nation‐ wide in their scale and the other two local。. One of the former is the just‐ mentioned Society,.

(3) M.UЁ No. though it is yet a sman groupo. 255. Another was organized in 1961 fOr. the purpose of cave explorations, caHed the Japanese Association for Caving,under the leadership of lⅥ [r. YAMAUTIo These two organizations have made several co_operative attempt to investigate Japanese caves. and have made many cOntributions to the knowledge of Japanese speleology.. In C)ctober 1959,when the Akiyoshi_dai Science Museun■. was founded. by the town of Shth6_ch6, Japan had a speleological museum for the flrst tiine.. It is 10cated On the Akiyoshi_dai, the karst topography of. which is the mOst beautiful limestone landscape in Japan and from. which we can gO dOwn by lift intO Akiyoshi_do. Since then the Museum has contributed much tO the advancement Of speleological knowledge of the Akiyoshi area. In 1961, the Museum established an underground laboratory fOr the purpose Of making physiological and ecological observations and experiments On cave anilnals. グ π sグ ′ %. The laboratory is set up in a small cave caHed KOmori_ana,4)abOut 200. meters 10ng, near the entrance tO Akiyoshi_do.. It cOnsists of three. rooms equipped with water supply and electric lights.. InvestigatiOns of anilnal and neolithic human remains have made a good start in sOme caves just in recent years.. 2. The Nu]mber,DilYlensiOn and the Distribution Of Japanese Caves There are known to be in Japan sOme 700 caves, more than 500 0f which have been scientiflcally investigatedo. The majOrity of them are. smal1 0r very small and only a few Of then■ are of moderate size. There are no lluge caves like the Marninoth Cave in Kentuckyo About 500 are limestOne caves and pot_hOles, 20 1ava caves located around volcanoes, and anOther 20 are deve10ped in the rock fOrmations other. than limestone, such as a few smaH caves formed in the ■ssures of sedilnentary rock at the f00t Of Mto Arafunё are prepared fOr abOut 70 1imestOne caves。 4) ana means cave alike dO.. , Nagano prefectureo Maps.

(4) Speleology in Japan. 256. Ryaga‐ do. γ απCθ びppθ r Eη ι 450 nl夕. Fig。 1。. `ふ. Plan of Ryaga‐ d6, northeast of the city of ]K6chi in Shikoku。. (Adapted from a map prepared by Mr.H.YAMAUTI) The largest lilnestone cave in Japan iS Akka‐. dO in the northern part. Of the Kitakami mountain range in the TOhoku district, measuring a little more than 8 kilometers in total length. Following Akka‐ dO in dilnension, both Akiyoshi― dO in the Akiyoshi area in the western part. cf Honsht and Rytga‐ dO near the Paciflc coast of Shikoku attain a total length of about 2 kilometers respectivelyo. Akiyoshi― dO opens at the. sOuthern edge of the Akiyoshi‐ dai, which is a conspicuous karst pene‐.

(5) M.UЁ NO. RY06‐ D0. Fig. 2.. Vertical sectiOn Of ]Rya6‐ d6 in Shikoku.(COurtesy Of Mro H.YAMAUTI; the flgure has been prepared by Mr.T。 OKAMOTO in 1964).

(6) Speleo10gy in Japan. 258. plane of 130 km2 in area and 200-400 meters above sea levelo. ThiS. plateau has numerous(circa 10,000)swa110W‐ holes(Doline)on its surface, and has sOme 130 caves and pot‐ holes, as well as compliCated. systems of underground streamSo The hitherto known deepest pot‐. hOle. was Rytgo¨ nO‐ COirait_ana located on the Akiyoshi‐ daio lt is a shaft Of 95 meterS in its vertical depth.. It Was named in honour of a French‐. man Dro Ho CoIFFAIT, Who had been a visiting speleologist in Japan in 1957 and co‐ operated with us in exploring caves in western HOnsht. and in ShikOku. In the early sumlner of 1964, however, YAMAUTI'S students explored a pot‐ hOle caned Ry色. the above‐ Inentioned one in depth.. 6‐. dO, which exceeds considerably. They were sucCessful in going. dOwn 216 meters to the bottom of that shaft from the upper entrance. It is located at Ono‐ ga_hara in the northwestern boundaries of]KOchi and Ehiine prefectures in Shikoku.. Lilnestone caves are scarCe in HOkkaidO and in the diStricts along the coast of the Sea of Japan,while they are numerOus in the districts along the PaciflC and in the southⅥ. restern parts of Japano. A great. number of caves are diStributed in Shikoku SOuth of its central mountain range, as well as in the Akiyoshi limestone area.. In the. former district, there are knOWn to be more than 70 caves, of which 60 were investigatedo. AInong therrl, Rytga‐ dO near the city of KOchi. is the most cOnspicuous in its complexity of passages with a great many cascades and pot― holes in the underground streams, and also in the richness Of its fauna. I haVe already mentioned the Akiyoshi area. Lava caVes are¬ wen deve10ped at the foot of Mt. Fuji(the Fuji vol‐. canO), and are also found in weStern Honsht and weStern Kytsht. sOme 40 caves are cOmmercialized ttor touriStS.. Stalactites and dO. stalagrrlites attractive to ViSitors are particularly beautiful in Akka―. in the T6hOku diStrict as WeH as in Ftren‐ dO near USuki in Kytsha. Rilnstone pools are wOnderfully developed in Akiyoshi‐. dO。.

(7) M.UЁ NO 3。. 259. The Fauna of Caves and Subterranean Waters lt is worth mentioning that Japanese caves are mostly small in scale;. nevertheless they have a rich and noticeable fauna. It is characterized by the predorrlinance of anilnals of prilnitive types as well as by the endenlicity of true cave‐ dweHers(trOglobionts5)). ActuaHy aH the troglo_. biontic species of anilnals hitherto found in Japanese caves are endeIIlic. to the lslands, and at the same tilne the majority of them have been regarded as representing old phylogenetic lines of respective groups. It is impossible to draw a sharp line between true aquatic cave animals. and true dwellers Of subterranean waters in the planes where there are nO caves.. This difnculty is an obstacle to the zoogeographic. presentation of the cave aquatic fauna as opposed tO cave terrestrial fauna. It is a noticeable fact that vertebrate cave d、 ア eHers are quite absent in Japanese caves, with an exception Of a marine gobioid ish Lπ α′ bπ s REGAN.. θ グ θgθ bグ %s. This white nsh smaller than 100 rrlin. in body length. with degenerated eyes has been found in a cave and a few wells near the sea coast in western Japan.. Arthropods are the most important. components of the Japanese cave fauna, as in European and AInerican. Fig. 3。. グ Lπ θ θgθ わグ %sα ′ わπs REGAN,found in a cave on the tiny island,. Daikonjima,in Shimane prefecture。 (After. TOMIYAMA,1936). 5)SOme American speleozoologists have used a term ′η gJθ bル θ for a true cave― dweller, but l am employing here the term ノ 知 g′ θみグ θη′ (Suttx biont denoting a living being)for it, in line with German research workers. “Naissance de la biosp6ologie". In an English sulrlinary of his recent article ηZα ノ 。ノ。sク θJθ θJ。 ,I(1964), (乃 C′ θ. 153-161),Co MOTAS haS applied troglobites to. a French terlm trog10bies(ScHINER,RACOVITZA). In hiS rグ ιγ♂θθgraったグ θ αグ σ々θ′ θgグ sθ 力ιγGγ πηご′ αgθ (1924),Richard HESSE,however, classifled cave animals in using the terms eucavale(rγ og′ θらグ θη),tyChOCavale(TrOglophilen) and xenocavale (TrOg10xenen). See alSO paratraph 4..

(8) 2δ. Speleology in Japan. θ. caveso. Some groups of aniinals belonging to this phylum have been. studied fairly weHo They are: trechine beetles, springtails, pseudo‐ scorpions, water‐. rnites, miHipedes, gammarids, cyclopoid and harpac‐. ticoid copepods, bathynenids and otherso. C)stracods and spiders have. not been sumciently studiedo Earthworims and natworlIIs have received. little attentione. Archiannelids are found in a few caves, but their. descriptions are not yet published.. YosII'S labOrious studies of springtails (Conemb。 la)for many years showed that there occurred in JapaneSe caves lnore than 100 species of this group,among which he deterΠ lined a large number of troglobionts. Jα , are of especial zoogeographic Some of thenl, such as the genus Sグ πθノ. interesto. ln this genus are known eleven species, six belonging to. Jα and the remaining ive to the subgenus Cθ the subgenus Sグ πθ′. θ θ θbrノ α .. YosII has pointed out that the Arnerican troglobiontic species of this. genus have developed from S.(S.Str。 )θ π″υおθ″ BR00K, While the グ (So Str.) λ∂漁′ Japanese representatives have been derived from S。 geography of cave¨ dweHing trechine SCHAEFFERo TaXOnomy and z∞. beetles(Harpalidae)haVe been So UЁ NO'S Chief work, on which he spent so much time since 1951 in exploring caves throughout the four main islands of Japan, as far as the Rytkya lslands and Taiwan. He described more than 60 species of troglobiontic trechids found in Japanese caves. His results enabled him to draw the conclusion that the speciation of that group of beetles in respective caves is remarkable.. This is best inustrated by his results obtained in Shikoku.. C)f the. 94 caves and pot‐ holes explored by hilnself, 25 have 20 troglobiontic. species belonging to four diferent genera. The genera(and Subgenera) change successively with the geographic distribution of caves from west to east in the following way: Rα 力απ″θθ力πs (yα 22η πガ蒻πs), θ 力πs(Mタ タθπθ‐ ルαηα′ ″θ 力πs(se str。 ),ISカ グ θ γθ ルαωα′ γθ θ乃%s(se Str.),ISカ グ Rα ルαπ′ ε 力%s。 γθ gα グθ πs(yπ αグθγgπ s), Rノ πg協 ごθπs(s. str.)and ttα ′ θ ttπ s), Rノ π γθ ′ α,η α occurs as a relict Besides these,a troglophilous species of r″ θθヵグ. in a cave at a high altitude in eastern Shikoku. Troglobionts are.

(9) M.UЁ NO. extremely. scarce in aquatic. 2δ ヱ. beetles on the earth.. In southwestern. Japan, however, there occur six species of bHnd dytiscids and phreato‐ dytids, the latter Of which is particularly interesting as they represent. an ancient stOck Of adephagous beetles and iH a gap between terrestrial and aquatic forms。. I.2″ 物 Fig.4.. ハ狗 γグ タ ηθノ θα ′カタα″グ θα s. uЁ NO, ♂ ,a blind aquatic beetle lθ. taken in the subterranean water at Himeji。. (After Shun‐. Ichi UttNo,1957). YAGINUMA haS described more than twenty species of cavernicolous spiderso. The majority of them are those found everywhere outside. caves; at least fOur or ive trog10biontic species are known, such as. OI, Ⅳ ♭s′ グθπs α力妙 θsカ グθ客 グS(UYEMURA), 0'レ θπs and Ce力 π″α′ 物9′ θグ YAGINUMA. Water_mites(Hydra_ carina)were studied by IMAMURA in recent yearso He described a Lθ ψ′ θπθ″. α力妙 θ磁 グ θ″gs. θh巾 ノグ グ YAGINUMA.

(10) Speleology in Japan. 2δ 2. great many new species found in subterranean waters and in caves。 s IMAMURA iS fOund 簿′ Among the cave water‐ mites, ルπごθθpsグs γノπgα θ夕 only in Rytga‐ dO and the other three species are endelllic to Akiyoshi‐. dO. I shan mention thenl in paragraph 4。. 魏 タ %s pα Jzゲ θ グαs′ ノgα θ αγ Fig.5.Two troglobiontic water‐ miteso A― びθ乃グ IMAMURA, ご ,ventral view.B一 』 石 グθθρsグ S'ソ π♂ π ηSグ S IMAMURA, ご ,Ventral. view.(After IMAMURA,1957,1959). It was an exciting happening when a rrlinute C)rustacea of a prilnitive. type was found by Y.. ルloRIMOTO. and Yo NIIuRA in the subterranean. water in the city of Hilnejio None of this group, Syncarida… Bathynel_. lacea, had ever been known in the regions outside Europe. Ⅵrith the. exception of the Malay peninsula. I and MoRIMOTO haVe been successful. in describing flfteen Japanese species 3α ′ ル Zげ π θ′ ″,Pα ″αbα ″げπθノ. belonging to three genera,. and a new genus A〃 θbα ′Zげ πθノル. established. by MORIMOTO and R/11uRA. All the species are troglobionts, some of which have peculiar features, such as keels in Pα M. UЁ NO and the large size exceeding 2 1nm. in」. タ カノπθノ zα ′ α ′ αγグ γαbα ′ αθ. Po gグ gク π″ θ α ⅣloRIMOTO.. Later l discussed the phylogenetic relation between the four genera γθ bα ′ ノ ブ レWhich is not distributed known in the world, including:「 力θ 々 ノπθ `“. ln Japan..

(11) M.UЁ No. Fig.6.. Pa夕. lα. ια′カノηθ′′ α gグ gα η′θα. 2δ θ. MORIMOTO,. in a driven well at the town of Dainan― Oita in Kyasha。. ,lateral view;taken. 。. ch6 sOuth of the city of. (After MoRIMOTO,1963). ITO has recorded more than t¬ Ⅳenty subterranean species of the cyclopoid Copepoda, most Of which are those found in surface waters.. Am[ong thelm,Spθ θcノ θゐpsノ θzθ θ′ 循グ S ITO is a lT10st noticeable species. HarpacticOid COpepoda are MIuRA'S chief wOrk, in which he. has. described mOre than ten subterranean specieso ln his recent monograph. on aquatic lsopoda, LIATSUMOTo gives a number of. subterranean. specles.. More than ten species of troglobiontic molluscs have been described. Both species Of the HydrObiidae, 4乃. well as. θsカ グ α πθπθグ KuRODA&HABE `り. ヵグ πθノ ル (J殷フ γグ α)α 力妙 θsカ グ θπsグ s βノ′. slowly flowing undergrOund streams.. aS KuRODA & HABE dWell in. The other troglobiontic snail. Cα υθ″夕 θηZθ ノ π′ ′ α 力πzπ %θ πsグ s. (SuzuKI)iS Senli_aquatic. Cavernicolous flat… worms(Turbellaria)are little knowne Two trogloxenOus triclads,D%gθ sグ α θα ノα′θπグ. lcHIKAWA&KAWAKATSU (it has long been identined with. Do gθ πθθθク物ル )and. P滋 ばθθα″. υグ グα (IJIMA&KABURAKI)are recOrded。 υグ ARNDT6)。 btained accurate statistics of the species of living anilnals. found in caves and subterranean waters in Gerlmany; but his meth_. od of cOmputation cannot be used in Japan at presento. HoⅥ. rever,. the numbers Of Japanese cave anilnals win further increase consider_ ablyo. Our knOwledge is still inadequate for certain zoogeographic. 6) ARNDT,Walther.Die Anzahl der bisher in Deutschland(Altreich)in Hё hlen und iln Grundwasser lebend angetroffenen メθttθ 力笏4g,1940,58-65。. ―笏πグ Kα′ ‐ Tierartene ■乙′ ″。H6ヵ ′ 夕 θ ι 省′.

(12) Speleology in Japan. 2δ 4. problems relating. to cave anilnalso. For instance, HOLDHAUS7)haS. pointed out that in Europe the northern limit of the distribution range of true cave anilnals other than beetles agrees to that of blind true. cave beetles. It is not certain whether such a phenomenon will be seen in Japan t00。. It will further be noted that there seem to exist. certain relationships between the JapaneSe and North AInerican cave faunas.. The faunal connection is particularly interesting in that the. mode of dispersal is markedly diFerent according to anilnal groups。 Certain groups occur only in the southeastern part of the United States. and in the Japanese lslands, while the others are spread over the northwestern areas of North AIIlerica and the northeastern corner of the Far East, including Japano. The former pattern is wen represented. by trechine beetles of the r″ θθヵθbノ θ22zπ S COmplex, and the latter by notopterans(GryHoblattoidea)Of the archaic orthopteran group. relations are exhibited also by the springtail group Sグ Cθ θ θθb″ ノα complex),chthOn五. Such. Jα ‐ θノ πθノ Jα (the Sグ π. d pseudoscorpions,and so ono This problem. must be solved by the close co‐ operation of Japanese and AInerican. speleobiologists. I expect and hope for a close contact between them for the promotion of speleology.. 4。. Cave Anilnal ColTlinunities No complete knowledge of anilnal cornlnunities has yet been obtained. for Japanese caves. From cave surveys we still know little about how their cave anilnal conllnunities are organizedo. The dimculties of such. surveys lie chieny in the actual labour involved in conecting in the dark, and deterrrlining the species of anilnals capturedo. The Spelaeo…. logical Society of Japan and the Japanese Association for{Caving have made several co‐ operative surveys of a number of caves, among which Akiyoshi‐ dO and RyOga‐ dO. anilnals living there.. have become best known with regard to the. I shall therefore take the former cave which is. 7) HOLDHAUS, K. Die europaische Hё hlenfauna in ihren Beziehungen zur ε α (Jena),コ (1932),1‐ 53. Eiszeit.Zθ θgθ θgπι ρカグ.

(13) M.UttNO. 265. most familiar to me, as an example to give an outline Of a cave anilnal. community in Japan.. p001s. Fig.7。. Plan Of Akiyoshi‐ d6.(Adapted from M。. OTA,1963). The anilnal cOmmunity in a cave dirers greatly in its structure from. any one of the epigeal communitieso. Every component of the former. is either dwelling sporadically everywhere in an eco10gicany favorable. situation or aggregated in particular places, viz. nlicrohabitatso. damp and Still. tt refer here tO all the anilnals in a heterogenous. assemblage in the cave under cOnsideration as the cOmmunity. ]Besides true cave_d、「eHers, trOglobionts(υ グ αθ απ′ θ, footnote 5), whiCh COuld not. live Outside a cave, there are usuaHy recognized twO mOre ecological.

(14) Speleology in Japan. 2δ δ. groups of cavernicoles, vize troglophiles and trogloxeneso Irlination of these two groups iS not always easy.. The deter‐. BOth cave anilnal. groups are as sirrlilar or the same as Outside species. Troglophiles are,. however, those whose populations haVe become capable of surviving their entire life cycle within the caves, while trogloxenes are found only frequently in caves. Beyond these we often meet u7ith aCcidentals which happened to be in caves. d6。. I shaH begin with the abiotic environment of Akiyoshi―. The. atmospheric temperatures in the depths of the cave are nearly constant. throughout the year, ranging from 12° -16° Co in January to 17°. -18° C.. in the nliddle of Julyo The annual amplitude is about 4° Co The area at a certain distance from the entrance receives the in■ uence of the. atmosphere outside the cave in wintero. Although it haS become. markedly dry in recent years, it iS always still wet because of an. underground. strearrl running throughout the cave after emerging. in the deepest part.. The annual amplitude of relative hullllidity is. extremely small(90-95フ 5).The temperatures of underground Waters do not exceed 18° C. in suΠ IIner and are higher than 13° . even in 1じ. winter. The water of bOth stream and pools is weakly alkaline, their pH values showing 8.0-8。 2.Owing to large calcium contents(52-70 ppm), the tOtal hardness of water gives large values(140-190 pprrl). The terrestrial corrllnunity of Akiyoshi‐ dO is composed of more than thirty‐. flve species of anilnals, the majority of which are regarded as. troglophiles or trogloxenes and have a lesser ecological signincance. Insects and spiders are the richest in the number of species, consisting of at least ten of the former group and twelve of the lattero Tγ π′ θ S. UЁ NO (S. Str.)タ ノ. θカグ α2π θ. iS a representative troglobiontic beetle8)and iS. the largest(7 mmo in body length)among the cave trechidso A camel πθ S力 θ″αzπ ″グFuRUKAWA,WhiCh ノ cricket rα θ ″θグ. is a common troglobiont. in the caves of southwestern Japan, is an important member, together 8) It waS alsO obtained at the bottom of two pot‐. hOles,. Ryago‐ n。 _coirait‐. ana. and Fasen‐ ana on the Akiyoshi‐ dai; in the forlmer it was found together with another troglobiont,Rα 々απJπ θ力2s(υ θZZι 解グ′″θ力2s)θ ′θグS.UЁ NO..

(15) M.UЁ NO. 267. with two troglophiles Dグ θs″ αππθ夕 η θ α ノα夕θπグ. BLATCHLEY and D.α タグθα′グS. BRUNNER,in the community because of the greater numbers of its in‐ dividualso. AInong the twelve species. of spiders,. six have become well. adapted to cave life, but there are only two troglobionts, Nes′ グ θπs αカグ γθ‐. s (UYEMURA) and πs の bα θ 鶴iグ グ YAGINUMA; the remaining. sカ グ ′ 徳グ θ. θ可. four. are. troglophileso. MiHipedes. (Chi10poda and I)iplopoda), the fOrmer. four and the latter six species, are also the important components of the conllnunity, though troglobiontse Fig.8.rγ cθ ヵグ αηzα (so str.)ρ Jπ ノ θ S.UЁ NO, ♂,taken in Nakao‐ d6. a志. 蝋. 器. Л. il,195の. there. are. no. AInong the chilopods,. three troglophiles Eク απθ″θ力θグπs θ′θグ γθ′″θ′ ルタ θ夕πS グ ルθααグ TAKA‐. MIYOSI,Sカ. 滋 s gπ ″″sは oc⇒ are KUWA and Oガ the common dweHers in this cave.. A nlinute amphibiOus snail, Cα υιγπ θη形′ια 力πzπ %θ πsグ s(SuzuKI), WhiCh iS 2 1Tllne high and be10nging to the Assiinineidae, is a representative. troglobiontic gastropod fOund On the wet waHs and rockse. C)f more. than ive species of springtails,Sグ πθノル (Cθ θbrノ α)α 力妙 θ〔 蒟グ α2η YosII is “ a typical troglobiont(υ グαθ απ′θ, paragraph 3).. There are found in this cave flve species of bats,9)an of which belong to the. insectivOrOus. grOup of the Chiroptera.. They are. trogloxenes which return to the outside of the cave periOdicaHy for feeding on insects,and yet they are of eco10gical signincance since their. 9) ]夕 πグρ′θ″πS sθ 乃%グ bθ 箱 グ /Jπ gグ πθsas HoDGSON, θノグ s zθ 知グαθノノJπ s 均 ′ TEMMINCK, Лう ′ θノグ S 2α ノ″ι紹 ″グ bθ π わグπ2s THoMAS, Rカ グ ηθ′ η ノクs ¢ クヵaS θθ夕 θθ夕 ηZZノ 2s TEMMINCK and R。 メθγγzzθ g究 グπ笏%π グ ρ夕θπ TEMMINCK. ZZイ.

(16) 268. Spele010gy in Japan. faeces(guanO)10)fOrm an important source. of food for cave. anilnals.. There are. sman accumulations of bats' dung falling ■oor. onto the. beneath the resting or. wintering site of the colonies, which have. decreased rapidly in number in recent yearso. No adequate information has yet. been obtained regarding Akiyoshi‐ dO一. the. bats. of. the size of their colonies,. their seasonal abundance and habits, and. what are the principal species forming heaps of excreta. ‐ α々グ ノθsカ グ. θπs Ncs′ グ. Fig。 9。. s(UYEMURA),a θ餐グ. trOg10‐. biontic. found in. spider. dealing with these matters。. In recent years we. Akiyoshi‐ d6。 (After Te YAGI‐. NUMA,1960;by perlnission of Hoikusha Publishing Co。 Ltd。. ,Osaka). cave.. 1 100k forward to Mr。. KuRAMOTO'S Careful work now in progress. ,. met occasionally. with two Species of rats, Rα ′′%s π ′′πs ″α″πs. (LINNAEUS)and Rα. ηθγυθgグ θπs. πs ″πs ttγ υθgグ θ. PALLAS, lWandering into the. They are accidentals WhiCh jOin the COminunity temporarily. with the coming of visitors. Regarding the f00d relation in the terres‐ trial community l will deal with that later one. The aquatic community is COmposed chieny of several species of Crustacea.. s ルθ々ππグ Psι παθθ″απgθ ηノ″ sカ グ. AKATSUKA`&KoMAI,Which is. widely distributed in the subterranean waters of central and Western Japan, iS the only troglobiOntic AInphipoda inhabiting the sha1low rilnstone poolso. Four surface Water species of the cyclopoid COpepoda,. sθ γγ%″ ′πs (FIscHER), %s(FIscHER)and ttSθ げ θノθps(So Str.)ル %θ ルα″′グ α′ θps ガ 物レ グ Pα ′ リッ θノ り s αノ蒻グπs θ′ し 」 θ θ′ ノ セ zθ γ. (JURINE),. Eπ. リ s ι θゐ ノ ソ τ. `Iソ. CLAUS Survive also in thoSe poolS as troglophiles or trogloXenese. There was 10). once found a population of a cladoceran Cθ. α γグ θααpヵ πグ. グθ:HUTCHINSON,G◆ E. Survey of conterrlporary knOWledge of biogeo‐ υグ JJ。 4夕ηθttθ αη. chemistry. 3. The biogeochemistry of vertebrate excretion. Bπ rso RQ′ 。二Jグ s′ .,New ハイι. York,96(1950),1-554.(Cave guano,pp.381‐. 461)..

(17) M.UЁ No. 2δ 9. (Oo Fo Mむ LLER).Water_■ lites are alsO the lmain components. gπ αグ″ απgπ ′ α. of the cOlmmunity;they areノ 物rθ πθ,SグS(■ 診χα″θπθ′sグ S)“滋%%α グIMAMURA, 0し 乃グ グレsttgα. γπS αル炒 θ磁 グ θ容 グ s. IMAMURA and Sθ ノααπθttb窃′″ α力炒 θsカ グθ. S. "扇 IMAMURA,an “ of which are trOglobionts and the latter two are endenlic. to Akiyoshi_doo A nlinute snai1 4ル グνθs力 ″α %θ πθグ eStablished by KuRODA. and HABE aS a new genus and a new species is a striking troglobiont inhabiting p∞ Is and slowly ■owing part of the streamo other aquatic invertebrates,Dπ ttsグα ′α′θπグθα. AInong the. ICHIKAWA&KAWAKATSU. is the Only triclad turbeHarian as a troglophile or trogloxene.. It is. worth mentioning that an archiannelid was obtained in the underground stream near the entrance of the caveo The food relation in the aquatic. cominunity is not known.. B. ノ 。 ノ. V ︲ I〃 プノ 、︱ ノ   ノ. r         . 燿︶. ‐ 11 『\. 日 丁︱にF上. A. ͡. lmm. 一 ノ. Fig。. 10. TwO cave‐ dwelling snails. A― ―Cα υθ. zα. θ 夕 2Zθ ′ α 力πzπ %θ ηsグ S ′. `η. (SUZUKI),shell and Operculum (Op)。. (Adapted from T. HABE,. 1941).B_4カ 妙θsカ グα %θπθグ KuRODA & HABE, two forms of shells.(Adapted from T.KuRODA&T.HABE,1954). During the past fOrty years since electric Hghts were instaHed in. the cave in 1925 fOr visitOrs, the natural cave envirOnment has gradually changed。. ヽ vithin the lilnits Of artincial lights, chlorophyH_. containing plants which fOund a chance to invade the cave have become capable of growing successfuny。. ` ■he recent improvement in the light. led sOme species of ferns and mosses to quicken their growth in the.

(18) Speleo10gy in Japan. 27θ. lighted area.. There are at present ive species of ferns and nine. species of mosses,11)as well as two Species of mushrooms, though the former two groups cannot fully grow up to a spore‐ forrrling stageo The ππ θ,“ グ ′ s 2%″ ノ ただαα PoIR. and Cノ γ′ principal species of ferns are fり ι″グ ππθグJ・ SM.Recently R/1re KANETSUNA haS fOund on the wret rock near 力 γ′ ιπ′ α α θθκ′ θπノ υグ. an artincial light eight kinds of diatoms, of Which Nα. was the principal specieso This diatonl is derived very likely from the. stream ■owing out frorrl the cave, because it makes up the flora there together with the other eighteen species of the same group. α,whiCh πθ α′ πル Vare′ グ タルθθπ′. Cθ. s θθπθ グ θ. is the mOSt abundant among them,was. Vas sucCessful αヽ θπ′ θπ′ αθ θπ′ incapable of entering the cave,whereas Nα υグ in flnding a new habitat under artiflcial light.. Some anilnalS COnstituting the conllnunity have become herbiVOrs in their food habit and feed on theSe green plantso. As is usual in moSt. caves, the priinary sourCes of f00d for anilnals in Akiyoshi‐ dO are organic debris transported into the cave by water and other agencies,. or introduced by man; as Well as bats' excretao. As a result Of the. growing of priinary producers Such as ferns, mosses or algae, the food. chain in thiS Cave, whiCh. had begun with the organic materials. mentioned above, has partly changed to Chlorophyll‐ containing plants in. the flrst instance.. Having enumerated twenty‐. tlⅣ. o Species of Cave. anilnals whiCh had been attracted to lights, KuRAMOT012)haS Suggested that certain trog10biOnts of ecological signincance in the COIIlinunity, απα, seem tO have πzπ πθ2簿 JS and Sグ πθJノ α α力″ ノθs力 ″ Jα ル such as Cα υθ″2zα ι222θ ノ The green plants growing in Akiyoshi‐ dO θ,Ser.B,2δ (1959), ′ 。 Sθ J.υ ″υ.ζ ノθ′ Cave,sOuthwestern Japan.ル 覺解.Cθ ′ ‐ グαグSε グ s乃 グ 」 BZ′ ′ ノθ , z4々 グ 315-322;HINO,I。 ,Plants growing in Shah6b6 cave.. 11) IWATSUKI,K.and S.UЁ NO,. .. 几グπs.,no.2(1963),26‐. 35(in Japanese,With English summary);HINO,I.,. ′ 。z4カ グ ノθsカ タグαグ Bryophytes fOund inside and outSide the Shah6do caveo Bπ ′ 56(in Japanese,With English summary). 2s.,no. 3(1964),49‐ イ Sθ グ . ハ shah6‐ dO is a synonyrrl fOr Akiyoshi‐ dO.. 12) KURAMOTO,T。. dO. ,Animals attraCted to the artincial lights in Akiyoshi‐ (In JapaneSe,with. ‐ 。ハイ2s.,no.2 (1963), 36-48。 グαグSθ グ Cave.B〃 J.4々 グ ノθsカ グ. English summary)..

(19) M.UЁ No. 271. changed their food habits tO dependence upon green plantso Regarding the nutritive relatiOn, Akiyoshi_do has now changed from a monotroph‐ ic cave13)。. f only aHochthOnous nutrition supply to an amphitrophic. cave,13) where f00ds are derived fronl both al10chthonous and autochthonous origin. We have no knowledge of cave bacteria and fungi which may play an important role in the food chaino. Micro‐. bio10gical studies will be necessary not only in Akiyoshi_do, but also in many cther caves in Japan. Such a development that has occurred near lights offers an interest_ ing ecological and physi010gical subjects tO be studiedo lt is, however, not always desirable to permit the existence of such conditions, which. will be unavOidable with the rapid increase of visitOrs to the cave. For the conservation of Akiyoshi_do, I think it win be necessary to eliminate such plants as much as possible in order to keep the natural conditions inside the cave, leaving certain typical lighted places for continuous ecc10gical observations。. 13) DuDICH,Ec,Die Klassinkation der Hё hlen auf biologische Grundlage.ル π″ ノ 助 力′ θ 2‐ 2η グ κα 箱″qθ ttθ 力。 ,Heft 3(1933),35‐ 43。 ..

(20) Speleo10gy in Japan. 272. Selected Bibliography of Japanese Speleozoology CHOPARD,Lo La distribution des Orthoplぬ res dans les grottes du JapOn.■ イ sん J, θ7(1963),101‐ 1090 DOTU,Y。 On the blind gobiOid iSh L7ε グθgθ ♭グzs αJbzι s REGAN.Zθ θ′.Mag。 (D6‐ zι. butsugaku Zasshi),72(1963),1‐. 5。. (In Japanese,with English summary).. GuRNEY,A.B.Further advanCes in the taxonolmy and diStribution of the Gry1lo‐ πgノ θπ,74(1961),67‐ 76. θ′ .Sθ θ.T石物sカ グ ・ ιグ blattidae(OrthOptera).Pη ε α new subgen。 ′ %θ ′ α (Cα υικ4α θ′ HABE,To On the recent specimen of Pα ′πググηθ′′. ). ηsグ s, with a list of known species of the Japanese cavernicolous たzz笏 笏θ πtts,12(1942),28‐ 32.(In Japanese,with English summary). molluscs.Tζ θ. IMAMURA,To Water‐ mites(Hydrachnellae and Porohalacaridae)from the subter‐ 工 Zθ θ ,12(1953),251-255。 ranean waters of Akiyoshi Karst,Japan.ノ αρ。 ―´ Subterranean Water‐ mites of the middle and southern Japan。 4π 功.ノ 。 ― 均 グ%θ bグ θ , 53(1957), 350‐ 391。 ― 一―――― Erste Mitteilung tiber Porohalacaridae aus unterirdischen Gewassern in 4 3π ttθ π,θ 5(1957a),53‐ 62. Japan。 4b力 .π αJ πγ ".yθ 乃 ― ―一 Some water‐ mites(Hydrachnellae)of Subterranean waters in Japane Aπ 461。 グ グ ′ %θ わ θ .,54(1958),447‐ ノ.力リ 一 ―― Check list of the troglobiontic Trombidiidae, Porohalacaridae and Hydra‐ .bグ θd婆 7ο gtt Sθ ε ・ Jα ραπ,21(1959),63-66。 chnellac of Japano B%′ ′ J。. J・. 0. 一―― ― Water‐ mites(HydraChnellae)Of Subterranean waters in Kanto district, θgグ α,1(1959a),426‐ 451. Japan.ス θαη′ δ8 44z。 , コ 。ノ 一―― Einige neue Arten von troglobiontischen WasserΠ lilben. Zθ θ′ ―一― (1962),252‐ 261.. lTO,T.CyClopoida copepodS Of Japanese subterranean waters. P″ ル θ′.び ″υ。 ,ゴ (1954),372‐ 416. ― 一 一 ――. western Japan.力. GroundWater copepods from south‐. (1957),1‐. Rθ 夕.FJS力・ 几αθ gJα リ グ筵 め グθ′θ. ,. コヱ. 28。. KOMATSU,T. Cave spiders Of Japan, their taxonorrly, chorology and eco10gy. 4『帆 乃πθJ.. . Eα s′ Asグ α, Sθ θ. KuRODA,T.&T.HABE◆. Osaka, Special Publ. 1961, 91 pp. 7z`ィ S,ヱ 9,for θ Troglobiontic aquatic snails from Japan.T'″. 1957 (1958), 183‐. 196。. MATSUMOTO,Ko TwO neW genera and a new subgenus of the family Asellidae π。 .ノ α ,θ 5(1962),162‐ 169。 夕θ of Japan.4π ηθJ.Zθθ′ ――一―― Studies on the subterranean isopods of Japan with notes on the well‐ water fauna of Japan.(Part I)。 Studies on the subterranean lsOpoda of ― グ.Sθ グ .,コ θ(1953),1‐ 77. ノ θttα b。 %ι θ 。αηπ ・ I:ι ク.助 乃ノθ Japano No。 1.Saク ρ′ ・ ク MIURA,Yo Subterranean harhacticoid copepOds frolm a driven well in Japan.Jα ノ。Zθ θ ,コ 4(1964), 133‐ 141. MIYOSI,Y。 もber iapaniSChe Diplopoden。 J。. Osaka, Special Publ. 1959。. o Eα s′ Asグ α ′ 。 Sθ θ ttη θ 五i+223pp.4‰αθ ,. (In JapaneSe).. ,H,Biol., MORIKAWA,Ko Cave pseudoscorpions of Japan(1).■ を解。Eカ ルπθ υ″υ。.

(21) M.UttN0. 27θ. 2(1956),271‐ 282. ― ― ―一. Cave pseudoscorpions of Japan(II).擁 %.Eカ グ解θ. υπグ υ。 ,H,BiOl.,2. (1957),357‐ 365. ―― ―. Systematic studies of Japanese pseudoscorpions。 II“. 2zo Eヵ ル2zθ 厖 夕. びηグ υ.,. B,4(1960),85‐ 172。. MORIMOTO,Y.Bathynellids of the Arnami group of the Ry色 ‐ Kya lslands(Syncarida, Bathynellacea).姥 %。 Cθ ノ ′ .Sθ グ υ.κ ノθJθ ,B,2δ (1959),269‐ 280。 .びπグ ―― Occurrence of a giant Pα 夕 わα′ヵノηθJJα in southeastern Kyasha, Japane ― lα. 4η ηθJ.Zθ θ′ .=嗜θη。,θ δ (1963),161‐ 166. MORIMOTO,Y。 & Yo MIuRA。 4′ ′θbα ノノηηθJノ α ′αρθηグθα gen.et spe nov。 , a new bathynellid from Japanc Pη θ .メ α 夕.4θ .Toyko,θ θ (1957),145¨ 148。. NoMuRA,S.Elmidae found in subterranean waters of Japan.4カ ルπ (KyOto),lθ (1961),1‐ 3。. TAKAKUWA,Y.Diplopoden aus Japan und ihr angrenzenden Gebietene vi+255 pp. Publ. by Jap. SOC. Promoto Sci。 , Tokyo, 1954。. with German summary)。. (In Japanese, partly. TAKASHIMA,H.&A.HAGA.A contribution towards the Japanese cave‐ dwelling species of the class Diplopoda.ル クsθ .Rθ 夕.yα %α Sカ グ πα's」 物sノ .Oγ π.& Zθ θ ′ 。 ,(1956),329‐ 343。 (In Japanese). UЁ NO,. Mo Notes on some subterranean isopods and amphipods of Japan.. 磁. 223。. Cθ ′ ′ θ 物 夕.υ ″υ。 。約 θノ ,B,θ (1927),355‐ 368。. ―― 一 The subterranean waters of Akiyoshi‐ 2 (1933), 91‐. 95。. dai and the fauna.t崎 .工 ニグ 筋,π θ′ .,. (In Japanese).. 一――― The Bathynellidae of Japan(synCarida‐ Bathynellacea)。 ζθ (1954), 519‐ 538。. 4π 力.二 均 湯つbグ θJ.,. - ObServations on the。 taxonolmy of the 138。 bathynenid genera.ノ 助 力々αグ 凌フυ″υ θ (1957),133‐ ,vI,ゴ A new suctorian parasitic to a Japanese bathynellid.乃 ―. .Fα θe Sε グ .. θgグ α,2θ リグ知bグ θ′. (With French r6sum6). UЁ No,So New cave‐ dwelling anchomenids of Japan.Oρ 2sθ .Eπ ″ 。 ,2θ (1955),56‐ 64. Studies on the Japanese Trechinac(V)(COleOptera,Harpalidae).ル %解 (1962),185‐. 187。. .. ― ― ―. .約 ′ θノ θ,B,22(1955a),35‐ 50.. Cθ ′ ′ .Sθグ .υ 〃υ. ]Blind aquatic beetles of Japan, with some accounts of the fauna Of Japanese subterranean waters.4π み.ノ 。均 グηιグ θ′ 。 ,5θ (1957),250‐ 296。 Studies on the Japanese Trechinae (VI)(COleOptera,Harpalidae).■ 姥2. Cθ 。Sθ グ .び ηグυ.κ ノθノθ,B,2イ (1957a),179‐ 218。 J′. 一― 一 The cave beetles from Akiyoshi‐ dai Karst and its vicinities. I. A new species of the genus rγcθ ヵグ α夕 ηα.II. こ ノθz π夕 ηグ %θ θ ノ ヵπs, a new group of the genus Rα 力αηノ 宅θ ttπ s.■姥解。Cθ J′ .Sθ グ .υ″υ.κ ソθ′θ,B,25(1958),39-48, 49‐. 61.. 一一―一― 「rhe cave trechids from the central part of the Chagoku district, Japane Io A new species of rγ cθ ヵグ αttzα from the Taishaku limestone area. II.. The geOgraphical races of rγ cθ んグα夕2oα. 27α グ ノθ力σノα夕. s.uЁ NO. M8夕 π.(Cθ ′′ ..

(22) Speleo10gy in Japan. 274. Sε グ .び ηJυ .約 θ′θ,B,25(1958a), 181‐ 184, 185‐ 197。 .び πグυ。 α α θ,zグ S.UЁ NO.MCπ o Cθ JJe Sθ グ 一一一一― Ditto.IIIo The group of 7kπ 乃グ `“ θ.B,2δ (1959),29-36。 Kノθ′ α (ColeOptera, Harpalidae)。 磁 解。Cθ J′・ ― ―― ― ― A synopsis Of the genus Kπ sπ 解グ Sθ グ .び ″υ.ζ ノθ′θ,B,27(1960),121-132. gα グ π A revision of the caVe treChidS of the genus ′ノπ ο S(COleOptera,. ―. θηo Trechinae)O Bπ .Nα ′グ J′. 。■こ%s.,Tokyo,8 Sθ グ. (1965),1‐ 16.. s′ 。 ツ物s. Ⅳa′ .〃 グ ,no.15(1962), YAGINUMA,T.Cave spiders in JapanO BZJ′ .Osα 々α 」 65‐ 77.(In JapaneSe,with English summary).. ー グαグ グθ sカ グ ′ Spiders frOnl lilnestone caves of Akiyoshi Plateau. Bπ ′. 4々 ノ s.,no.2(1963),49‐ 62.(In Japanese,with English summary)。 Sθ ブ .■んι ノ%,2θ (1954),62‐ 70. YOSH,Ro HOhlencollembolen Japans I.κ θηノ Zθ θ′。 .工 ,11(1956),609‐ 627. ψ ル 一 ― 一 H6hlencollembolen JapanS H・ hlencollembolen JapanSo Cθ π′γ。 わグθ′.二 αbo Xフ θ′θ ____ Monographie Zur Hё ― ―. び力υ。 ,no.3(1956a),109+22 ppe ―― ―― (Э n the geographical diStributiOn. of cavernicolous Collembola. Bπ. JJ.. π,2θ (1958),13‐ 17。 (In Japanese)。 gtt Sθ θ gθ θ .Jα クα bグ θ θ θわ句ρ Cθ θ πθ α‐ 一 一一一一 SOme ColleIIlbola frorn Okinawa caves, with notes on Sグ 34。 ・ グ グ α グ Sθ .ル物 グ グ θ sカ ,no.3(1964),25‐ 4々 〃 。 ノ cOmplex of Japan.Bπ J′. s。.

(23) 275 UЁ NO:Speleology.. 1.. PLATE. ア a group grOup of rilnstOne pools in A_kiyoshi― dO. traフ , a “Hyakumai― zara",. (COurtesy of Mr.T.KuRAMOTo). The K6mori_ana UndergrOund Laboratory attached tO the Akiyoshi― Science Museum。 (COurtesy of Mr.T.KuRAMoTo). dai.

(24)

Fig。  1。   Plan of Ryaga‐ d6, northeast of the city of ]K6chi in Shikoku。
Fig。  10. TwO cave‐ dwelling snails. A― ― Cα υθ

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