Transactions and Proceedings of the Palaeontological Society of …100S... · 2015-12-21 ·...

Transactions and Proceedings

of the

Palaeontological Society of Japan

New Series No. 100 Supplement

A Concise History of Palaeontology in Japan

Edited by

MATSUMOTO, T.,

T. HAMADA, H. UJIIE and Y. TAKAYANAGI

(English editor: R. FUSEJIMA)

Palaeontological Society of Japan

June, 1976

Preface

At the annual meeting on January 25th, 1975, our Society decided to publish "A Concise History of Palaeontology in Japan" as No. 100 supplement of the Transactions and Proceedings of the Palaeontological Society of japan, New Series, to commemorate the continued activity of the Society since its foundation in 1935. This has been materialized by cooperation of a number of contributors under the editorship of a committee specially organized for this purpose. We are very happy to present here this commemorative volume. On behalf of the Society we thank many of the members who have given valuable suggestions and/or financial support to enable us to publish this concise but memorial number 100s, and Honorary President Dr. Teiichi KOBAYASHI and other contributors to the following chapters.

Apart from a short note entitled " List of fossil subgenera, genera and higher taxa proposed through Japanese publications" compiled by T. HAN AI and K. KONISHI, which appeared as an appendix to Geology of japan (TAKAI, MATSUMOTO and TORIYAMA, ed., 1963), the present publication is the first attempt of a comprehensive review of the history of palaeontology in Japan written in a Western language. In view of the great advancement in palaeontology and related sciences in recent years, the need of retrospects and prospects is keenly felt in each field as well as in the interdisciplinary domain. The present work has been intended to meet these requirements, although each article is done as concisely as possible. Most of the references cited in the articles are found in " Bibliography of Japanese Palaeontology and Related Sciences, 1941-50; 1951-1960; 1961-1975" published in Palaeontological Society of japan, Special Papers, No. 1 (1951), No. 9 (1962), and forthcoming issue (1977). For brevity we omitted duplicate listing. It is our eager hope that the present publication may serve for improvement and advancement in future palaeontological studies at home and abroad.

Editors

\


Introduction to the History of Palaeontology in Japan

Teiichi KOBAYASHI*

Education and Research: It was the early quarter of last century that LAMARCK restricted the term fossil to organic remain. Until the last quarter of the century, however, little has been known of fossils in Japan. H. Th. GEYLER was the first to publish Fossile Pflanzen aus ]uraformation japans, Palaeontographica, Band XXIV, 1877, denominating Podozamites reinii for a land plant in favour of J.J. REIN who procured it at the foot of Volcano Hakusan, Province of Kaga (Ishikawa Prefecture), 1875.

A Survey of Fossils from japan Illustrated in Classical !yfonographs, 1963 consists of reproduced illustrations of ten articles by ten pioneers which were published in Europe and their revised identification. GEYLER was followed by NAUMANN, SCHWAGER, MOJSISOVICS, NATHORST, BRAUNS, TEMPERE, YOKOYAMA, NEUMAYR and JIMBO. YOKOYAMA's article was the seventh, 1890 and JIMBo's the last, 1894, both on Cretaceous fossils. The others were SCHWAGER's fusulinids, MOJSISOVICs' Triassic ammonites and pelecypods, NEUMAYR's Jurassic corals, brachipods and molluscs in addition to YOKOYAMA's foraminifers, NATHORST's Mesozoic and Caenozoic land plants and NAUMANN's elephants. Thus various aspects of Upper Palaeozoic and later fossils were already sketched in these articles. It was just about the dawn of Palaeontology in Japan. _

It was the fifteenth century that Leonardo DA VINCI challenged the tradition of

* University of Tokyo, Professor Emeritus.

the Noachian deluge and Lusus Naturae. Since the Renaissance Natural History progressed in Europe steadily. In the Orient Chu HSI1 correctly understood the origin of petrified shells some 300 years before DA VINCI, but the matter was not probed deeper to the bottom. In China, however, was a long history of Pentsao2 ,

probably from the first century. Including the knowledge on stone medicines Pentsao was imported to Japan through Korea in the fifth century and made great advances in diverse trends since the seventeenth century. Sekitei KIUCHI (1724-1808) recorded in Unkonshi3 (1773, '79, 1801) various rocks and minerals including coin stone (crinoid), stone turf (coral), nails of a long nosed goblin4

(shark's teeth) and so forth. This book may be comparable to Lapidarium Sinicum5 by H.T. CHANG", 1921. Such collecting and appr~ciation of curios bore no scientific value by themselves, but the ardent curiosity of the so-called Roseki (stone-trifling) school7 combined with the serious efforts of Dutch scholars and others of Oriental Classics during the age of national isolation prepared fertile ground for west!;!rn seeds of palaeontology and other sciences to be transported and soon naturalized in Japan.

In 1877 when GEYLER's paper was published in Germany, the University of Tokyo was established and palaeontology was begun to be taught. Edmund NAUMANN was the first professor of geology

1 *;a 2 *lj!! a ~t&;t 4 ~E. Ez, ::R~IiJJT\E 5 E~ "*~iU 7 WEi*

Trans. Pro c. Palaeont. Soc. Japan, N. S., (lOOs), 1976

2 MATSUMOTO, T. et al., edit.:

and his chair was succeeded by Toyokichi HARADA in 1885 through David BRAUNS and Carl Christian GOTTSCHE. BRAUNS, Geology of the Environs of Tokio was printed in the i'vfemoirs of Science Department, Tokio Daigalw1 (University), No. 4, 1881. YOKOYAMA graduated from the University in 1882 and was appointed professor of palaeontology in 1889 when his jurassic Plants from Kaga, Hida and Echizen was published through jour. Col!. Sci. Imp. Univ., vol. 3, part 1. In 1882 the Geological Survey of Japan was established and in 1893 the Geological Society of Tokyo (later Japan) founded and published its journal. Thus Tokyo was the center of education and research in geology and palaeontology in the Meiji era (1869-1912).

Products of this era were studies on Mesozoic land plants and Jurassic marine molluscs by YOKOYAMA, fossil echinoids by YciSHIWARA (later TOKUNAGA), Desmostylus by YOSHIWARA and IWASAKI, Cretaceous ammonites by Y ABE, Tertiary foraminifers, corals, bryozoans, gastropods and Lithothamnium by NISHIWADA, NEWTON and HOLLAND, Neogene pteropods by YAM AKA W A and [W ASAKI and so forth. Y ABE's Mesozoic plants and Gigantopteris from Korea, 1905 and 1908 and YOKOYAMA's Mesozoic and Palaeozoic plants from China, 1906 and 1908 were early contributions to the continental palaeontology. YoKOYAMA's Climatic Changes in japan since the Pliocene Epoch, 1911 gave rise to a moot .discussion in Japan and abroad.

Matajiro YOKOYAMA (1860-1942) devoted much of his time from 1910 to 1932 to the studies on Tertiary and later molluscs from various areas from Karafto (Sakhalin) to Taiwan (Formosa) which greatly urged the advancement of Caeno-

zoic biostratigraphy of the festoon islands. Because this series of his papers is such an indispensable reference to Tertiary students, its taxonomic revision was carried out by MAKIY AMA, T AKI and OYAMA in the light of recent conchology in Special Papers of the Palaeontological Society of Japan, Nos. 2 & 6, 1954 & 1959 and OYAMA's re-revision on the type specimens from the Kanto area in Spec. Papers, No. 17, 1973. Most of Japanese palaeontologists are, needless to say, either his direct pupils or indirect ones through Y ABE, MAKIY AMA, 0ZA W A and others. Therefore, to commemorate this progenitor the Palaeontological Society of Japan prepared its society medal with YOKOYAMA's relief and conferred it first to Y ABE in the Twenty-fifth Anniversary of the Society.

YOKOYAMA ·was an excellent writer of popular science. His ·books propagated palaeontology aud related sciences widely in Japan. Many of technical terms and taxonomic names of his version into Chinese characters are internationally used in the Orient. Early in 1891 he published Japanese Text-book of Palaeontology1 in a concise form. Later he wrote two standard hand-books with titles of Prehistory of the World 2 (Historical Geology), 1918 and Elements of Palaeontology3, 1920. This work was succeeded by 13 palaeontologists and Palaeontology in 2 volumes was published in 1954 and 1957. Now Palaeontology is in the way of complete revision by many authors and 3 of 4 volumes appeared already in 1973-75. In short, it is not too' emphasizing that the education and research of Japanese palaeontologists were originated from YOKOYAMA.

In 1912 the second geological institute was established at the Tohoku Imperial

A Concise History of Palaeuntology in Japan 3

University. It was split into two institutes in 1922, namely Institute of Geology and Palaeontology and Institute of Mineralogy, Petrology and Economic Geology. For the establishment of the geological institute Y ABE came to Sendai and conducted education and research with his colleagues forcefully and results were published in Science Reports of the institute. H. MATSUMOTO published many papers on fossil mammals of Japan and China in the early volumes (1915-30) of this publication. It was an interesting coincidence that this and Memoires Serv. geol. l' Indochine, Hanoi were two equally gigantic palaeontological publications in Eastern Asia both started in 1912.

Hisakatsu Y ABE (1878-1969) is the other important founder who set Japanese palaeontology on the firm basis. His works in the science are so numerous and extensive that his papers are countless and cover nearly every branch such as foraminifers, corals, brachiopods, ammonites, other molluscs, mammals, rockforming algae, land plants and so forth. Among them, however, he devoted particularly to the ammonites, corals and foraminifers. His contributions to the geology of Japan and adjacent areas are not the less important. Since 1925 he had particular interest in the coral-reef problem which has led him to visit many south sea islands. It is of crucial importance that he educated many leading geologists and palaeontologists on one side, while on the other he supported scientific societies and activities. In fact, the Palaeontological Society of Japan would not have been founded so early without his effort. He was not only elected honorary member of many domestic and foreign scientific societies but also awarded Cultural Merits (1952) and Cultural Medal (1953) from the Department of the Imperial Household.

Until 1945 three more geological institutes were established in the Imperial Universities at Kyoto (1921), Sapporo (1930) and Fukuoka (1939) in addition to a few others in other universities. Beside these institute publications a large amount of palaeontological papers were published in Proceedings of Imperial Academy (1912-, later Proc. japan Acad.) and japanese journal of Geology and Geography which was primarily (1922) published from the National Research Council (1920-49) but from the Science Council of Japan from 1949. It is a great pity that the latter organization abandoned in 1975 the publishing of this journal ( 45 vols.) together with some other Japanese Journals, all internationally well known.

Members of the Geological Society of Japan were about 200 in 1910, but increased to 900 in 1940. In 1935 the Palaeontological Society of japan was established as a section of the Geological Society. Papers of the Palaeontological Society were published in its section of the other's journal and these reprints were compiled under the title of Transactions and Proceedings of the Palaeontological Society of japan. The number of its members was 283 at the beginning among which were several biologists and about 20 foreign members. They are now 523 in total, about 10 percents being foreign members. In 1957 the Society became independent from the Geological Society. Prior to this New Series of Transactions and Proceedings and Special Papers began to be published in 1951. Subsequently in 1960 a journal in Japanese named Kaseki or Fossils was added to them.

From 1949 Japan entered the age of many universities, and palaeontology has been taught widely in the course of geology or earth sciences. In 1962 a


palaeontological laboratory was set up in the National Science Museum, Tokyo which was primarily the Education Ministry Museum, 1872.

As outlined above, thanks to European scholars palaeontology as a science of Natural History was imported to Japan in the last quarter of last century and studies on the fossils from the continent were already commenced at the beginning of this century. The Geological Survey of Korea was instituted at Seoul in 1918, Shanghai Science Institute in 1926, Geological Institute of Taihoku (Taipei) University in 1928, Central Museum at Changchung, Manchuria and Tropical Industry Institute at Palau, both in 1938.

On the continent RICHTHOFEN travelled China extensively for four years from 1868. His China in 5 volumes was its outcome which was the source book of the primary importance for the geology of Eastern Asia. Volumes 4 and 5 of China contain various fossils described by specialists. Subsequently, Loczy (1877-1880), WILLIS and BLACKWELDER (1903), J. Coggin BROWN (1907-11), J. DEPRAT (1909) and others repeated geological reconnaissances in China and their collections greatly amplified our knowledge. The Tokyo Geographic Society undertook such a reconnaissance in central and south China in 1911-1916. Y ABE and HAYASAKA's Palaeontology of Southern China, Reports of Geographic Research in China, vo!. 3, 1920 include3 many Palaeozoic fossils. In 1933 the First Scientific Expedition to Manchuria was carried out under the leadership of S. TOKUNAGA. Its reports included 4 volumes in geology and palaeontology. During the following ten years or so such scientific reconnaissances were repeated to Manchuria, Inner Mongolia, North China and other areas by Japanese

parties. Among the palaeontological results

achieved on the continental side are the studies on the Cambrian and Ordovician faunas, Cathaysian flora (KAWASAKI and KoN'No), non-marine Mesozoic fossils (K. SuzuKI et a!.), discoveries of Silurian and Devonian fossils (K. OZAKI et a!.), late Pleistocene Dalainor skull with early Mesolithic implements (R. ENDO, 1945), exploration of contemporaneous remains at Kushantun near Harbin and so forth. On the Pacific side those are on larger and smaller foraminifers, living and fossil corals, a deep well in Kita Daito-jima (North Borodino Island) and others.

In these 30 years our palaeontological studies were made on the North Pacific side (T. MATSUMOTO, KANNO et a!.) and further in the Andean region on one side and in Southeast Asia, Near and Middle East and Africa on the other. Their results are so far partly published. One of them is Geology and Palaeontology of Southeast Asia in 15 volumes (1962-) which clarified Cambrian and later biostratigraphy of the zone from Thailand to the Philippines through the Malayan Peninsula and Borneo. Amud (Neanderthal) man (H. SUZUKI and TAKA!, 1970) was found in Palestine by Tokyo University Expedition toW estern Asia. 1961-74. NAKAZAWA's party found Permian relic brachiopods and pelecypods in the lowest Skytic fauna in a continuous Permo-Triassic section in Kashmir. Incidentally, S. MATSUSHITA (1926) discovered a productid in the Skytic Kurotaki limestone, Shikoku Island, long ago.

Here a brief mention is added as to the international co-operation through IPU and other organizations. Japan has participated early in the originating meeting of the International Palaeontological Union which was held at Cosmos Club on the occasion of the XVI

A Concise History of Palaeontology in Japan 5

International Geological Congress at Washington, D.C., 1933. Since then our palaeontologists and society always supported IPU (now IPA) in various ways. Its Asian Filial (now Regional branch) meeting was held at Tokyo in 1965. Japanese palaeontologists often cooperated with the Pacific (formely PanPacific) Science Congress in its standing committees on coral reef, correlation, etc. or symposia, such as a symposium on Tertiary Correlation and Climatic Changes in the Pacific (ASANO, HAT AI), 1965, and with the International Commission on Stratigraphy, lUGS, formerly IGC, as active members, in the compilation of Lexique Stratigraphique International in part of Asia, vol. 3 and Oceania, vol. 4 (HANZA w A) for example. So are they at present with International Geological Correlation Programme (IGCP). They participated in many international conferences and colloquia in geology and biology insofar as they were related to palaeontology.

I have written above the growth of the palaeontological circle in Japan and its national and international activities in three ages of Tokyo, several universities and many universities, although the later the age, the shorter the statement. Now, shall we see the aspects of the fossil biota of Japan and her surroundings and the achievements of our palaeontologists through these ages.

Palaeontography or Descriptive Palaeontology: As usual in natural history, palaeontology of Japan started with descriptive work. Catalogue of Type Specimens of Fossils in japan, 1961 records type specimens of 5856 species of plants and animals from Japan, Korea, and Manchuria beside some other areas which were described mostly by japaneses but partly by foreigners from 1888 to Sep-

tember, 1960. In its 360 pages 187 (90, 74, 22), 54 (28, 20), 38 (25, 11), 26, 24 (14, 9), 12 and 9 pages are occupied respectively by Mollusca (Pelecypoda, Gastropoda, Cephalopoda), Protozoa (Smaller and larger Foraminifera), Planta (Vasculares, Non-Vasculares), Coelenterata, Arthropoda (Trilobita, Crustacea), Brachiopoda and Vertebrata. The remainder includes Echinodermata, Bryozoa, Porifera, Annelida and Problematica. Roughly a half of the species belongs to Mollusca (a quarter to Pelecypoda), one-seventh to Protozoa and one-tenth to Planta. In other words, more than three-quarters of them are objects of palaeoconchology, micropalaeontology and palaeobotany. This is an aspect of the above fossils. HAY AMI's A Systematic Survey of the Mesozoic Bivalvia from japan, 1975 contains 867 species which consist of 236 Triassic species, 247 Jurassic ones and 284 Cretaceous ones.

Mop.ographic works were already published on fossil woods (SHIMAKURA, 1936), Proboscidea (MAKIY AMA, 1938), Cenozoic brachiopods (HAT AI, 1940), Tertiary smaller foraminifers (ASANO, 1950-52), Mesozoic corals (EGUCHI, 1951), Upper Palaeozoic calcareous algae (R. ENDO, 1951-61), Coralliaceae (ISHIJIMA, 1954), Permian and Carboniferous corals (MINATO, 1955), Cretaceous foraminifers (TAKA Y ANAGI, 1960), Palaeogene Mollusca (OYAMA and MIZUNO, 1960), Jurassic ammonites (SATO, 1962), echinoids (NISHIYAMA, 1966, 68), Lower Cretaceous pelecypods (HAY AMI, 1965-66), Silurian trilobites (KOBAYASHI and HAMADA, 1975) and so forth. In addition there are Illustrated Catalogue of East-Asiatic Fossil Plants by OISHI, 1950, leones of Fossil Plants from japanese Islands by S. ENDO, 1955, and Index Fossils of japan by SRIKAMA, 1964.

List of Fossil Subgenera, Genera and


Higher Taxa proposed through japanese Publications in Geology of japan, 1963 comprises 1249 taxa in total, including 1079 genera and subgenera (A) and 170 subfamilies and higher taxa (B). In A group Mollusca occupies almost a half, followed by Arthropoda (1/5), Coelenterata, Protozoa, Brachiopoda, Vertebrata and others. B group consists of 90 taxa of Arthropoda, 41 of Mollusca, 11 of Echinodermata and smaller numbers of Coelenterata, Brachiopoda and Vertebrata. It is noteworthy that Mollusca is, as in the above species number, first in A group and second in B group, but Arthropoda is different. It is fifth in species number, but second in A group and first in B group. This means that Arthropoda becomes proportionally larger in higher taxa in the list whereas Mollusca is largest or almost so in all of the three groups.

HANAI and KONISHI (1963) outlined systematic palaeontology in Japan. in a very concise but well condensed form. Histological studies on Araucarioxylon Tankoense, Cyathocaulis naktongensis, Cycadeoidella japonica, and many other fossil woods were done by STOPES, Fum, OGURA and other botanists. Metasequoia MrKI, 1941 is a fossil genus still surviving in China ; Tingiostachya KoN'No, 1929 is a cone with which Tingiales was erected in the Articulatae ; Nilssoniaceae was proposed with Nilssonocladus by Ta. KIMURA and SEKIDO, 1975. Propilina is a Monoplacophoran genus. Metanothosaurus is an aquatic Triassic reptile; Manchrodon a Pantotherian mammal from the Jurassic of Manchuria (YABE and SHrKAMA, 1938); Nipponosaurus an Upper Cretaceous ornithopod from Sakhalin (NAGAO, 1936). They are only a few genera among rare or conspicuous fossils. lJIRI's histological study on Desmostylus' teeth is a nice piece of work.

A copious Pleistocene fauna was described from fissure deposits of Kuzuu, Tochigi Prefecture (SHIKAMA, 1949). Pleistocene Nipponoanthropus akashiensis was collected from near Akashi, Hyogo Pref. (NAORA, 1931). Additional human bones of Palaeolithic age are known from Mikkabi, Shizuoka Pref., Ushikawa. Aichi Pref. and Iejima, Okinawa Pref. (TAKA! and SUZUKI).

Palaeontography of the japanese fossil biota is at present fairly well advanced as to the leading groups, but much remains to be done on some minor groups or ill-preserved ones. Sponges, belemnites and crinoids belong to such groups. Studies on the Cirripedia, insects and fishes are recently improving. Neither graptolites nor cystoid is as yet uncovered in Japan.

Classification, Phylogeny, Ontogeny and Evolution: OzAwA's classification and zonation of the Fusulinidae, 1925 were later developed by Y ABE, HANZA W A, FUJIMOTO, TORIY AMA, KANMERA, and others. The family revision was carried out further on the Polymorphidae (OzAWA, 1931), Dipteridaceae (OISHI and HuzrOKA, 1936), Halysitidae (HAMADA, 1957), Waagenophyllidae (MINATO and KATO, 1965), Pteriidae (ICHIKAwA, 1958), Myophoriidae (NAKAZAWA, TAMURA, et al.), Trigoniidae (YEHARA, NAKANO, et al.), Turritellidae (KOTAKA, 1959), Tertiary Pectinidae (MASUDA, 1962), Cytherideidae (HAN AI, 1959), Lycopteridae (TAKA!, 1943) and other families of Tertiary foraminifers and molluscs, Cretaceous ammonites, older Palaeozoic trilobites and so forth. Y ABE and SUGIYAMA have shown a phyletic rejuvenescence of the Stromatoporoidea in the Jurassic Torinosu sea.

In pointing out the deviation of the Volborthellida from the principal lineage of the Cephalopoda, the incipient diver-


gence of Ordovician uncoiled nautiloids was traced back to late Cambrian Plectronoceras whose siphuncle was no.t holochoanoidal (1935, 37) .. The final divergence of late Mesozoic ammonites was, on the other hand, explicitly illustrated in Japan as the result of prolonged research from YOKOYAMA to MATSUMOTO through Y ABE, SHIMIZU, and others. Nipponites is a classical example of the most aberrant form indicating a terminal specialization.

While K. SAITO (1936) discovered a spheric protoconch of Hyolithes globiger, ventro- and centrosiphonate types of apical ends were distinguished among Ordovician nautiloids (1937). HANAI (1953) discovered a new structure called primordial rostrum behind the protoconch of Neohibolites miyakoensis. The regular apical septation was found in LO\ver Cambrian Helcionella and Hampilia.

Because the Proparia and Hypoparia are polyphyletic, combination of biocharacters was emphasized for the natural classification of the Trilobita. The Agnostida, Redlichida, Corynexochida and Ptychopariida were recognized as independent primary orders, because they indicate four Lower Cambrian original stocks. The ontogeny was clarified of Blackwelderia quadrata (R. ENDO, 1935) and Redlichia chinensis (KOBAYASHI and F. KATO, 1951). HANAI (1951) has shown parthenogenesis of " Cypris " subtriangularis and sexual dimorphism of Cypridea subvaldensis. HANZA WA (1964) discussed polymorphogenesis of the Lepidocyclinidae and Miogypsinidae.

MAKIYAMA (1924, 41) exemplified the :volution of minor scale with Neogene Umbonium and Siphonalia. Recently HAY AMI (1975) unraveled population variation of Pliocene-Pleistocene Cryptopecten vesiculosus. Prior to this the time-relation

of W AAGEN's mutation to DE VRIES' mutation (i.e. saltation) was elucidated by means of zone-time (1944, 45). The Glyptagnostus hemera was pointed out to be the oldest world instant in the Phanerozoic. eon (1949).

Biostratigraphy, Palaeogeography and Palaeoecology: As indicated by SMITH's second law of stratigraphy the rock sequence of the Japanese islands was extended back with the discoveries of Lower Carboniferous, Devonian and Silurian fossils respectively by I. HAYASAKA (1922), Y ABE and M. NODA (1933) and ONUKI (1937). The oldest so far known is the copious Silurian fauna described by SUGIYAMA (1940) and HAMADA (1958). Ordovician ellesmereoceroids were discovered in New Guinea (KOBAYASHI and BURTON, 1971).

In the early days the geological age of fo3sils, accordingly fossil beds, was determined by direct comparison with European fossils in the classical sequence. Therefore YEHARA's division of Cretaceous rocks by means of Japanese trigoniae. (1923) bears extraordinary importance in the sense of natural division of rock sequence with reference to vertical and horizontal distribution of indigenous fossils. The bio3tratigraphic classification was considerably improved by MATSUMOTO's intensive studies on Cretaceous rocks and contained ammonites and Inoceramus in which the latter NAGAO cooperated. The Cretaceous system would be most advanced in fossil zonation in Japan. It is divided into six series and twelve stages. Many life zones were distinguished in three groups of ammonites, namely, the Desmocerataceae, (1), Acanthocerataceae, Hoplitaceae, etc., (2) and Heteromorpha and Lytocerataceae (3) beside inocerami. The whole sequence of the system was correlated with the


classical standard in Europe and the Cretaceous sequences of the Tethyan and Pacific areas in detail.

Of the provincialization it is noteworthy that the Phanerozoic biosphere began with three provinces, namely the Olenellian, Redlichian and intermediate provinces where the last connected the two others (1971, 1972). The biostratigraphy in Eastern Asia bears great importance for the interprovincial correlation and the system boundary, the Ozarkian problem for example (1934).

The history of land plants is well documented in Japan and her adjacence from Devonian (TACHIBANA, 1950) onward. The northeastern limit of the Cathaysian phytogeographic province is marked by the Kaishantun flora at the southeastern corner of Manchuria (KONNO, 1969). Its expansion into the Japanese islands was confirmed by ASAMA in the Abukuma and Kitakami mountains. It was thoroughly proven in the Upper Triassic paralic sequence by marine faunas that the so-called Rhaeto-Liassic flora by OISHI and others appeared in Japan already in the Carnic epoch (1939). Subsequently Carnic and Noric plants and their fructifications were described from West Japan (KoN'No). The climatic fluctuation in Japan and Eastern Asia during the Mesozoic periods was analyzed with land plants by the change of their composition. CHANEY, T ANAl and other Japanese palaeobotanists studied Tertiary plants in many areas in the islands from 1958 to 1972 and considerably clarified the floral change during the Tertiary period and the problem on the ancestors and descents of the Miocene forest in the Pacific area.

Four suite3 of non-marine Mesozoic faunas were distinguished in Eastern Asia through studies on molluscs, fishes and others. One of them was the Con-

chostraca. As the result of its extensive revision it was found that the center of its distribution in Eurasia shifted from the west in the Palaeozoic to the east in the Mesozoic era in accordance with continental growth by orogenies (1954, 75). The extensive distribution of Trigonioides and its allies in the Cretaceous formations in Asia and similar shells in North Africa and western North America casts a question on their origin and migration. The land connection of the islands with the continent was discussed with mammals by Y ABE (1929) and others.

As done by SHUTO, CHINZEI and many others, palaeoecological and biostratonomical studies were carried out with various materials in different ways to decipher the bio-, thanato- and fossilhistory which fossils and fossil associations bear. An interpretation was given on diatom thanatocoenosis from the North Pacific (KANAYA and KOIZUMI, 1966). Some interesting studies were done on trace fossils and problematica. Deformation of Triassic ammonites was discussed by MAKIY AMA (1942) and defossilization in Radiolarian rocks by KoBAYASHI and To. KIMURA (1944).

Palynology (S. TOKUNAGA, K. TAKAHASH!, et al.) and nannoplankton studies became very active in last two or three decades. ASANO, TAKAYANAGI, UJIIE, et al. demonstrated the stratigraphic significance of planktonic foraminifers. Conodont studies by leo and others from 1963 have shown much of Triassic sediments included within the Upper Palaeozoic formations of previous reference.

C. TSUBOI and HIRATA (1935) reported the increasing tendency, from Recent to Miocene, of the angle of the long axis of aragonite fibres with the C-axis of aragonite crystals in the shell of Glycymeris yessoensis. In recent years new techniques, biochemical, radiometric,


palaeomagnetic and mathematical, or by means of a scanning electron microscope, X-ray microanalyser, an electronic computer or else were applied to p::llaeontological and related investigations in Japan as in other countries. Ultramicrostructures of various fossils were observed and their palaeobiochemical analyses made (HONJO, AKIYAMA, I. KOBAYASHI, et al.); biometric studies done with molluscs, larger foraminifers and others (OBATA, HAYAMI, UJIIE, MATSUMARU). Colloquia were thrice repeated recently on isotopes in hard tissues of fossils (KONISHI, TAKAYANAGI).

Finally, a mention is added that KOBAYASHI (1946) elucidated the relationship of documents to synthesis in palaeontology and historical geology. Subsequently IJIRI (1949) discussed method and cognition in palaeontology.

For palaeontological literature of Japan the reader is referred to Bibliography of japanese Palaeontology and Related Sciences, 1941-50 and 1951-60 compiled by R. ENDO and TAKA! and printed in the Palaeontological Society of Japan, Special Papers, Number 1 (1951) and 9 (1962), respectively. Bibliography of japanese Palaeontology, 1961-75 is now in preparation. He can trace them back further through Titles and Abstracts, in japanese journal of Gzology and Geography, 45 vols. 1922-75 and Index to the journal, vols. 1-20 (1955), 21-25 (1960), 26-30 (1961), 31-35 (1968), 36-41 (1972) and 42-45 (1975) and with Bibliography of History of japanese Earth Sciences, compiled by the Committee for History of japanese Earth Sciences, Tokyo Geographical Society and to be published through journal of Geography, Tokyo.

Summary: Retrospect and Prospect: The above history of palaeontology in Japan is no more than a general view. Palaeon-

tology was proposed almost simultaneously by Ducrotary DE 8LAINVILLE and Fischer VON W ALDHEIM (1834). This history of education and research is what happened in japan in the later two-thirds of the time length since the proposal. It is divisible into three ages and the research achievements into three fields, systematic, palaeobiological and biostratigraphical. In this article more weight was laid in the older age and in the achievements on the international bearing rather than local interest. The reader will, however, find in the succeeding articles detailed informations including those which were omitted because of page restriction. Regrettably I feel that allocated time was too short for such a topic, although I have endeavored to meet the society's request.

Palaeontography is the prerequisite for palaeobiology. The systematic description of the fossil biota in Japan advanced slowly at the beginning, but the Meiji palaeontology is astonishing for me in that it was about a dozen of years to absorb the western palaeontology so much that Meiji palaeontologists could develop their science principally by themselves thereafter. It was considerably accelerated in the Showa era (1926-) through the Taisho intermezzo (1912-26). The basic work of this kind was done also on the continental side as well as on the south sea side until1945 and later in Southeast Asia and further beyond and in some Pacific areas. In consequence the achievements bore a perspective aspect. Two notable points in the Showa palaeontology were that fossils were touch stones for the existing palaeontology and that biostratigraphic facts were oriented from the global standpoint for high synthesis. These viewpoints combined have enabled any fundamental and international contributions for Showa palaeontologists.

to MATSUMOTO, T. et al., edit.:

The purpose of our work was at the beginning to establish the stratigraphic sequence of Japan and then that of adjacent areas. This trend of research required the accuracy of zonation on one side and the interprovincial correlation on the other. At the same time it developed into palaeoecology, biostratonomy, p::tlaeogeography, p.1laeoclimatology and so forth.

Fossils are essential documents of evolution. Therefore the evolution and the vicissitude of the biosphere in the Phanerozoic eon are two principal subjects of palaeontology. Phylogeny and ontogeny of some fossils were clarified as much as suitable materials were available. Some new structures were discovered and new classifications of certain families and higher categories proposed. The migration, dispersal, provincialization, rise and fall of plants and animals

were investigated on certain groups with reference to geographic and climatic changes and other factors.

These studies shared the advancement of palaeontology and related sciences in part of the Asian and Pacific sides with the effect that it reduced the unbalance in our knowledge between these sides and the European-Atlantic side to some extent. Therefore it may be said that it advanced an important step toward the creation of a complete world statue of palaeontology. The method of palaeontology, however, has been conventional during several decades, although there were minor devices and some ameriolations. Recently new techniques are affording previously unavailable data. Therefore, future development is promising, as traditional palaeontology will tie up with new palaeontology in researches and syntheses.

Paleontology and Society of Japan

Hiroshi U JIIE*

Japanese culture of today was developed, for the greater part, during the Edo period that lasted 265 years until the Meiji era began, mainly owing to the national isolation without disturbances by war. The culture bred up under such circumstances attached great importance to subjective contemplation and sense of beauty. The society that fostered the culture rejected logicality as a mere argumentative or unrefined way of thinking. Thus the Japanese society of the Edo period lacked the ground for

* Department of Paleontology, National Science Museum, Tokyo.

presentation of objective views of things or systematization of the results of observation, and so the development of the science of natural history was much hampered. The Edo people esteemed natural objects highly only when those were beautiful or extraordinary, and the hobby of collecting rare objects prevailed. Among the collectors known as " stonelovers " was Sekitei KIUCHI who made corect sketches of stones and presented his views on their origin (1773-1801). But KIUCHI was an exceptional case of amateur collectors.

On the other hand, some branches of learning attained a high level, as repre-

Trans. Proc. Palaeont. Soc. Japan, N.S., (lOOs), 1976


sented by Wasan (the mtive mathematics of Japan) which was developed for practical p'.lrposes. However, practic:1l use of technical le1rning was circumscribed within certain professions which were again restricted by the hereditary system. Accordingly, the accumulated knowledge was transmitted only from father to son or fro:n teacher to disciple, and was seldom allowed to spread beyond the closed circle. Without sufficient communication true science could not be developed.

Mining was a matter of primary concern of the Edo government, and it accelerated practical application of technical knowledge. Toward the end of the Edo period when the country was prepared to be opened to foreign intercourse, there rose a trend to absorb Western learning. Genpo MITSUKURI, who was on the staff of the Foreign Documents Survey under the government administration, wrote a number of books introducing Western civilization to Japan. His works included " Chishitsu Bensho " (Dialectic Geology) (1861) and two other handbooks of geology. Nevertheless, he contributed little to the development of geological sciences in Japan, because most of his books were in manuscript, which prevented wide circulation, and his knowledge did not go beyond translation of literatures.

During this period, foreigners began to come to Japan. Some of them collected animal and plant specimens and fossils to be studied in their own countries, but the results of their study were seemingly insignificant from the world-wide view since people's interest was aroused in the fact that the specimens came from " Gipang ", the land of dreams.

In 1868 the Meiji government was born, and the nation-wide modernization started. It was strongly felt that Japan at that time was behindhand in modern

industries. The waves of colonialism of the Western Powers were invading the coasts of the Oriental countries. Under such circumstances, exploitation of underground resources was a matter of urgent necessity for Jap:ln. To meet the national demands, the government invited experts from foreign countries, such as B. S. LYMAN from America in 1872 and E. NAUMANN from Germany in 1875. These foreigners fulfilled their duties in surveying underground resources, but at the same time they endeavored to bring up their successors among Japanese. Particul3.rly notable were the achievements of NAUMANN, who taught geology at the Tokyo Imperial University since 1877 when the Institute of Geology was established in the Faculty of Science. He also helped founding in 1878 the Geology Section (later Geological Survey of Japan) under the Bureau of Geography belonging to the Ministry of Home Affairs.

During the early part of the Meiji era, foreign teachers and technical experts took the lead in geology and paleontology, but after 1894 the graduates of the Tokyo Imperial University and the persons who finished their studies abroad began to take phce of these foreigners. It was fortunate th:1t the Japanese forerunners in geology and paleontology made efforts in raising the level of learning and educating students, without seeking fame or prominent post. In the meantime, Institute of Geology was set up in seven univenities, from the Tohoku Imperial University (1911) to the Hiroshima University of Science and Literature (1943). Well-trained paleontologists were assigned to these institutes to teach paleontology. It was a most steadfact development for those days, if we recollect the postwar period when rapidly increased universities have suf-


fered from shortage of good teachers until quite recently.

Along with the progress of modernization of nation!!.! constitution, Japan began to extend her territory over the . neighboring regions, and this inevitably gave rise to geological survey organs and corresponding agencies, as exemplified by the Geology Section (hter Manchoukuo Geological Survey) that was established in 1907 in the South Manchurian Railway Company after the termination of the Russo-Japanese War (1905), and the Geological Survey set up in 1918 in the Government-General of Korea after the Japan-Korea amalgamation was enacted (1910).

Apart from the political problems, geological and paleontological know:Iedge gained from these regions was of great value, especially with regard to Mesozoic and Paleozoic eras. To the Japanese geologists and paleontologists the data on the Chinese continent were indispensable. The results of their research activities, keeping out of politics, are classics still useful to this day. In comparison with the above-mentioned period, the later times that were marked with the outbreak of the China Incident in 1937 and the outset of the Pacific War in 1941 produced little noticeable results in spite of the fact that a great many geologists and. paleontologists traveled all over the Cl1jnese continent and Southeast Asia, carrying out field work and collecting specimens to be brought back to Japan for study. The poor results can be attributed largely to the national policy of those days demanding urgent exploitation of underground resources, as well as to the worsened economy due to the long-continued wartime, but the scholars and the academic society may have been partially responsible for the consequences.

In 1945 Japan had her first experience of a lost battle, and during the succeeding several years the country was to3sed about by inflation and a surge of the social system reform.

The Association of Democratic Scientists was established in 1946, and the next year saw the inauguration of the Association for Geological Collaboration which thereafter took the lead in innovation of geologists circles in Japan. The Association's activities contributed greatly to equalization of opportunities and to destruction of sectionalism among research bodies, but on the other hand they incurred antagonism with competent but conservative groups. With the passage of the last thirty years, however, the left-wingers and the right-wingers are seemingly advancing toward reunion.

After the several years of postwar disorder wore away, publishing circumstances improved. A number of textbooks began to be published, and academic journals were reissued or newly published. The Transactions and Proceedings of the Palaeontological Society of Japan was separated from the Journal of the Geological Society of Japan in 1951 to make a start as New Series.

Economy of Japan became prosperous owing to the special procurement boom caused by the Korean war that broke out in 1950. In 1959 the balance of the Bank of Japan notes exceeded 1,000,000,000,000 yen, and Japan entered into the age of high growth of economy. In the same year, the liberalization of dollar exchange came into effect, and in 1960 foreign trade and international exchange were liberalized. These circumstances were greatly favorable to international conferences and cooperative researches that were beginning to take place. Japanese geologists and paleontologists to be dispatched for, or participate in, inter-


national scientific activities abroad increased year after year, and overseas survey parties were sent out. From 1963, the Ministry of Education compiled a budget to defray funds for overseas researches. Thus, the " Paleontological Studies of Southeast Asia" that had been financed by nongovernmental sources came to receive government support since 1964. The purposes of these overseas survey parties were entirely academic, differing from the wartime precedents, and in most cases the survey work was performed in collaboration with the native organs, and so considerable results have been gained and reported. It must be noted, however, an upsurge of nationalism of the country to be surveyed is striking these days, and it is often the case that dispatch of easygoing survey teams is rejected under the pretext of underground resources protection or nature conservation. Review of the history of Japanese paleontology may be of a help toward true international cooperative survey and study.

Among the geological sciences in Japan, paleontology has been taking the most academic course. With the introduction of various kinds of analytical equipment, microscope and computer in the 1960's, Japanese paleontology seemed to advance toward paleobiology and paleophysiology, but in broad perspective the effect of social changes on paleontology was remarkable. In order to make the effect more positive, we need to extend such fields as biostratigraphy and paleoecology, and develop practical application of them. Paleontology, originated as a branch of geology, cannot be expected to cultivate and develop its basic fields without a pipeline of application. In this connection, we must look straight at the fact that the Geological Survey of Japan, staffed with more than 500 geologists,

comprises no section of paleontology, and that no micropaleontologist is on the staff of any marine research institutions in this country.

Supplement : Brief Review of the Transactions and Proceedings (New Series) of the Palaeontological Society of Japan.

The Palaeontological Society of Japan, that had started in 1935 as a section of the Geological Society of Japan and had been publishing the papers of its members in limited pages of the Journal of the latter Society, came to issue a journal of its own, entitled Transactions and Proceedings, New Series (quarterly), from 1951 on. This was due to the stabilized Japanese economy after the war, as well as to the increase in its members (514). Animated publication of academic journals was a universal phenomenon of those days; for example, Bulletin of the British Museum (Natural History), Geology (the first issue in 1949), Neues Jahrbuch fiir Geologie und PaHi.ontologie (1950), Vaprosy Paleontologii (1950), Revista Italiana di Paleontologia e Stratigratia (1952), Senckenbergiana Lethaea (1954) and Micropaleontology (1955), which are enjoying worldwide fame now, made their first appearance one after another in those days.

For the last 25 years, publication of the journal was continued through the efforts of many scholars and senior researchers including those who had already passed away, and the journal has come to be known internationally, as evidenced by the fact that about onethird of the subscribers are persons and institutions of foreign nationals. At this point of time it may be worthwhile to review the contents of the journal to provide for future growth (refer to Fig. 1).

Fig. 1 shows the changes in the relative


YEARS .j

OJMMULATIVE PERCENT NO. of ARTICLES

Fig. 1.

frequency of the articles printed in the journal during the period from 1951 to 1975, with two years as the time unit (which is also the unit used in numbering and indexing text pages and plates). According to the figure, fusulinids (denoted by Fu), Paleozoic molluscs (pM) and Mesozoic molluscs (mM) excluding ammonites hold constant, though small, ratios. On the other hand, Cenozoic molluscs (eM) and larger fossil plants (PI, mostly leaves), which used to account for large proportions, are decreasing lately; this may be explained by that the methods of study of these branches which had experienced a long history of

research are too classical for the articles to be duly evaluated. The same tendency is noticed throughout the world. Vertebrates (V) also show a similar trend but it is probably due to the limited occurrence of fossil vertebrates in Japan and to the scantiness of vertebrate paleontologists. The ratios of corals (C), brachiopods (B) and trilobites (T) have been rather constant, although the number of articles is small, on account of limited occurrence of specimens. More sporadic ·are the articles on arthropods (A) excluding trilobites.

Now, our interest is aroused in the fields whose vicisitudes reflect the epi-


sodes in the history of science. In the field of Foraminifera (Fo), Kiyoshi ASANO and a few others contributed the articles in the early stage, and then there came a period of a remarkable decline between 1956 and 1960. After that, younger researchers began to write articles from various points of view until 1970, but the young power turned apparently inactive when it became plain that for the foraminiferal study up-to-date international information on planktonics and new technique for benthonics are requisite. Also in the field of Bryozoa (Bry), study of which was started in 1957, articles were very few between 1967 and 1969, although it was due to personal reasons on the part of researchers who were no more than two in number.

On ammonites (Am), the lack of articles from 1958 to 1962 is ascribed to private circumstances of Tatsuro MATS UMOTO who was the major contributor, but from 1962 on MATSUMOTO and his students, and also Yuji BANDO who dealt with Triassic ammonites, published their results in succession. Especially the biometric approach attempted by Ikuwo OBATA and Kazushige TANABE is worthy of notice.

Study of conodonts (Cn) became active since MULLER visited japan and disclosed their occurrence, but few articles have appeared in the journal, whereas in foreign countries monographs are being published successively. As conodonts are very important for the study of Japanese late Paleozoic to Triassic Systems, much expectations are laid on further progress in this field.

Microfossil plants (MP1, excluding pol-

len and spore) were represented until 1960 by the works of Haruo OKUNO who studied microstructures of fossil diatoms by means of electron microscope. From 1969 after a conspicuous gap, articles began to appear but the subjects are entirely different from the previous ones and moved to the biostratigraphy.

One of the greatest missions of paleontology would be contribution to the comprehensive earth sciences through biostratigraphic studies. With regard to this point, articles on micropaleontology dealing with pollen, spore (Po) and other microfossils would naturally require wide space for listing the specimens that are found in great numbers. Other fields than micropaleontology also need enough space for geologic maps to indicate occurrence in time and space of the materials dealt with. In this connection, the journal has hitherto failed to offer sufficient space for reporting the results of studies. Since 1963, the limited number of pages for each article was increased from 12 to 24, and the number of plate from 1 to 2. As a result, the number of articles to be printed was reduced (see the numerals on the right column in Fig. 1), but it is a fact that the subjects became manifold and the level of articles rose. A prize for the best paper published in the Transcations is also stimulating contributors since 1966.

Constant efforts are desired for the improvement of the journal so that it can print not only the representative articles on paleontology in japan but also a number of excellent reports from abroad.


Paleozoic Microfossils

Hisayoshi !Go*

Fusulinaceans : The first study of Japanese Paleozoic microfossils goes as far back as the introduction of Fusulina japo. nica by W. GUMBEL (1874), which started the dawning of the research activities by foreigners such as C. SCHWAGER (1883) and J. DEPRA T (1914). Hisakatsu Y ABE took the initiative in the study of fusulinaceans also, and reported his results in 1899, 1902, 1903 and 1906. The proposal of the genus Neoschwagerina in 1903 came from his very far-sightedness. lchiro HAY AS AKA (1924) published his study on the Omi Limestone and described fusulinaceans contained therein. Roughly simultaneously with HAY ASAKA's work, distinguished•· reports by Yoshiaki OzAwA (1925, 1927; -etc.) on the Akiyoshi and Akasaka Limestones were published and attracted universal attention. Fusulinaceans of Indochina were studied by COLAN! (1924) and those of North China by LEE (1927), and the both works were published in bulky volumes. Their results, along with the series of OzAwA's work, made a long step forward in the researches on fusulinaceans of Eastern Asia in the 1920's.

In the 1930's the researches by DUNBAR of North America and RAUSERCHERNOUSSOVA of USSR made progress, while Shoshiro HANZA w A, Haruyoshi FUJIMOTO [ =HuziMOTO] and others of Japan were actively working. Y ABE and HANZAWA (1932) announced their new view on the classification of fusulinaceans and proposed the genus Pseudodoliolina.

* Institute of Geoscience, The University of Tsukuba.

FUJIMOTO (1936) published an excellent report on the fossils from the Kanto mountains. A paper by HANZAWA (1938) on Nipponitella created a sensation among the fusulinacean researchers of the world. His work (HANZA w A, 1938) on Pseudoschwagerina and Zellia was important as well, and initiated the discussion of the Carboniferous-Permian boundary problem.

As the time lapsed into the 1940's, Ryiizo ToRrY AMA took part in the research on fusulinaceans and described specimens from the Yasuba Conglomerate and other formations (TORIY AMA, 1942, etc.). HANZA WA (1941) discussed the Carboniferous-Permian boundary in Japan, Korea and Manchuria, attaching importance to the absence of the Triticites zone. His description of Parafusulina yabei is, though brief, one of the eminent papers.

Soon after the termination of W or!d War II, study of fusulinaceans in Japan began to flourish, having been reinforced with young and energetic researchers. The postwar researches were based on a detailed stratigraphic study in the field, entirely different from the method of prewar time. Especially noticeable among the numerous researches are the following:

TORIY AMA (1954, 1958, etc.) made an exhaustive study of the Akiyoshi Limestone, and added new information to the knowledge of the geologic structure of the region since the works of OzAwA and Teiichi KOBA Y ASH!. At the same time, he established detailed fossil zones, among which the Profusulinella zone was



recognized for the first time in Japan. in 1958, TORIY AMA's work on fusulinaceans was published in a bulky book of 264 pages and 48 plates. In the latter half of the 1960's TORIY AMA and his collaborators worked on fusulinaceans of Thailand. In 1967, TORIYAMA summarized the fusulinacean zones of Japan.

From the 1950's to the first half of the 1960's, a large number of distinguished reports by Kametoshi KANMERA were published. He is known for his scrupulous descriptions of fusulinaceans and conscientious preparation of plates. Since 1952 he published, in succession, the results of his studies of the Yayamadake Limestone, the Kuma Formation and the Kosaki Formation. These reports, including the paper published in 1959, on the evolution of Neoschwagerininae; are notable as they raised new problems on the Upper Paleozoic stratigraphy and fusulinacean zones of Japan. The Fusulina, Triticites and Pseudoschwagerina zones, as well as the Kuriki Series and the Hikawa Series, all proposed by KANMERA, still serve as the standard stratigraphic units. The study of the Sakamotozawa Limestone by KANMERA and MIKAMI (1965) was remarkable in that it revealed details of litho-facies variation of the limestone and the occurrence of fusulinaceans.

Roughly contemporaneous with KANMERA, Ken'ichi ISHII made a new epoch in the study of the Japanese Carboniferous fusulinaceans on the basis of his researches on the Itatorigawa Group of Shikoku. He scrutinized the postwar papers by Soviet researchers, which were gradually becoming available in those days, and he re-examined genus Fusulina and revived genus Beedeina. His papers published during a period from 1955 to 1962 brought forward some new ideas about the evolution of fusulinaceans.

During the same period, there were a series of works by Rokuro MORIKAwA who developed the "sump" method for examining specimens, free from the microscopic observation of thin sections, and made a unique classification (1955, 1958, etc.). It is regrettable that he died before the completion of his study with "Solidgraph" (1962), into which all his energy was put forth.

Hisayoshi IGO (1956, 1957, etc.) published the result of his study of the Ichinotani Group, which revealed the almost perfect succession of the Millerella, Profusulinella, Fusulinella, Fusulina, Triticites and Pseudoschwagerina zones. Later, IGO (1964, 1965, etc.) studied the Nyukawa Group and considered the relationship between the litho-facies and the occurrence of fusulinaceans. In recent years, he has been proceeding with the study of fusulinaceans of Thailand and Malaysia (IGo, 1972, etc.).

Yasuo NoGAMI (1961) published the result of his detailed study of fusulinaceans from Atetsu-dai. In 1965 he reexamined the original specimens previously described by SCHWAGER. With the successive studies of fusulinaceans in the Taishaku-dai Limeatone (Kimiyoshi SADA, 1961, 1963, etc.), the foasil faunas of limestones in the Inner Zone of Southwest Japan were clarified to a considerable extent. Reports by Susumu HONJO (1959, 1960) and Masao MINATO and HoN-· JO (1959, 1965) are worthy of notice in that they introduced new viewpoints in the study of fusulinaceans. They used enlarged photographs of Neoschwagerina, etc. for the plates, and examined the details of development of septula and other features.

During the period of 1950's to 1960's more results were published ; important ones among them were the works of Michihiro KAWANO (1961) on Yamaguchi

18 MATSUMQTO,. T. r?t al., edit.:

Prefecture, Manabu KOBA Y ASH! (1957) on lbukiyama, Mosaburo KANUMA (1958-1960) on the Mino mountainland, Shigeo SAKAGUCHI (1963) on Tamba, Kazumi SuY ARI (1961, 1962) on Tokushima Prefecture. Kunihiro lSH!ZAKI (1962) on Kochi and Ehime Prefectures, Takeshi CHISAKA (1960, etc.) on the Kitakami mountainland, Yoshinari T AKAOKA (1960) on the Kanto mountainland, Shigema KAWADA (1954. etc.) on the Omi Limestone, and Masafumi MURATA (1961) on Akiyoshi-dai. Other noticeable results were; the consideration on individual growth of Pseudoschwagerina by Saburo AKAGI (1958), the report by Atsushi ISHII and Hajime TAKAHASHI (1960) on Paradoxiella, the study of the southern Kanto mountainland by SAKAGAMI (1957, etc.), the report on Mesoschubertella by SAKAGAMI and KANUMA, the report on Vetbeekina by Tomomitsu SuGr (1960), the o;:currence of Fusulina from Hida and other studies by Kenji KoNISHI (1952, etc.) and the study of the lse-Shima district by Nobuo Y AMAGIW A (1956, etc.).

In the meantime, the scholar3 who had been active in the prewar day3 published some important studies and theories by making effective use of their wide experience, and thus gave an impetus to young researchers. Y ABE (1949) was the first to point out the existence of the Millerella zone, and on many occasions, from 1964 to 1966, he discussed the problem of Lepidolina and the Carboniferous-Permian boundary. HANZA WA (1949) proposed Acervoschwagerina, and in 1950 and 1954 he published interesting papers on Eoverbeekina and Afghanella. In 1963, collaborating with MuRATA, he published a paper as a warning against too much circumstantial classification of fossils, citing instances of Neoschwagerina, etc. FUJIMOTO assisted by !GO proposed the genus Hidaella in 1955, and

summarized the fusulinacean zones of the Carboniferous System (1958).

The study of fusulinaceans in japan seemed to pass the peak since the end of 1960's, and TORIY AMA, KANMERA, !GO and Ken'ichi ISHII moved the focus of their researches into Southeast Asia. Nevertheless, Tomowo OzAWA (1967, 1970, 1975, etc.) continued to publish the results of his outstanding work and established a new way of •.learning supplemented by statistics. His study of Pseudofusulinella and his comments on the evolution of Verbeekinoidea are worthy of special mention, as are his excellent work on Lepidolina multiseptata and its evolutional change. Of late, the achievements of younger researchers, including the works of Dong Ryong CHOI (1975, etc.) on the Kitakami mountainland, of Fumio KOBAY ASH! (1973) on the Nagaiwa Formation, and of Kozo WATANABE (1974) on the Omi Limestone, haye been published and great hopes are placed on their future efforts. The fusulinacean researches in Japan are far from complete yet, as there remain many fields to be investigated and not a few problems concerning paleoecology and evolution are still unsolved. Smaller foraminifers : Late Paleozoic limestones of japan contain abundant smaller foraminifers associated with fusulinaceans. Nevertheless, the study of these foraminifers was very slow to start. 0ZA WA (1925, 1927) and FUJIMOTO (1936) were the pioneers in this branch of science. Yuji OKIMURA studied the Atetsu Limestone and other Carboniferous limestones in various parts of the country (1958, 1965, etc.) and brought forth a lot of new information. His earlier theory on Paleozoic chronology, though not sufficiently circumspect, should be rated high in respect that he set his hand to a difficult task. Future development of his work is expected.


Radiolarians: The study of Paleozoic radiolarians was commenced by FuJIMOTO (1938) with an intention to clarify the geologic age of cherts which are commonly found in the Mesozoic and Paleozoic Systems of Japan. His study developed into the discussion of the age of the Sanbagawa crystalline schist. He maintained that Paleozoic radiolarians can be discriminated from Mesozoic ones by the percentage of Cyrtoidea. His view was opposed by Teiichi KOBA Y ASH! and Toshio KIMURA (1944) and KIMURA (1944), and this gave rise to an active controversy in the academic world. Koichiro ICHIKAwA (1950) worked on the radiolarians of the southern Kanto mountainland, and his age-determination has proved correct in view of the present knowledge of conodonts. Researches on radiolarians of both Mesozoic and Paleozoic are very important and future progress is expected. Ostracods : Researches on Paleozoic ostracods are much behindhand in Japan.

There are the work of Takashi HAMADA (1959) and the serial studies by Kunihiro IsHIZAKI (1963, etc.). Further development of this branch is anticipated as the material for study is fairly abundant. Conodonts : The history of researches on conodonts is still very young in Japan. It began with the work of IGO and Toshio KOIKE (1963) and Shingo HAYASHI (1963). Afterward, with the progress of the study of Carboniferous conodonts by !Go and KOIKE (1964, etc.) and KOIKE (1967), it was revealed that considerable amounts of Triassic conodonts are contained in the limestones and cherts which had been regarded as Paleozoic (KOIKE et al., 1971, etc.). This revelation brought forth not a few topics for discussion of the PermianTriassic boundary in Japan. At present, conodonts are known to occur in all of the Paleozoic formations of Japan, ranging from Silurian to Permian, and their stratigraphic significance is highly esteemed.

A Brief History of Post-Paleozoic Micropaleontology

Yokichi TAKAYANAGI*

In Japan, study of Post-Paleozoic microfossils was begun at the end of the 19th century. The first papers on foraminifers and diatoms were published in 1890 or thereabout, but those were succeeded by little research activities until the early part of the 20th century when Hisakatsu Y ABE commenced his study of foraminifers. Micropaleontological researches in Japan today are active and manifold, but

* Institute of Geology and Paleontology; Faculty of Science, Tohoku University, Sendai.

the history of study of great many groups of miCrofossils is still short. It may be safely said that micropaleontology in Japan developed centering around researches on foraminifers, and so in this article the history of foraminiferal study will be introduced by dividing · it into several stages which also cover the historical progresses in various other branches.

The initial stage of micropaleontology in the 19th centuray was opened in the form of the study of Japanese specimens

Trans. Proc. Palaeont. Soc. Japan, N. S., (lOOs), 1976


in Europe, as exemplified by the work of J. BRUN and J. TEMPERE (1889) who studied diatoms, and Matajiro YoKOYAMA's study (1890) of foraminifers, etc. Though the source of the original material of the diatom study by BRUN and TEMPERE remains still obscure, their work helped to let the Japanese diatoms known to the world. The circumstances of the early period of foraminiferal study were described by Y ABE in his memoirs (1953). YOKOYAMA's report (1890) on the foraminifers from the Paleogene of Hokkaido (though he first assigned the age to Cretaceous) was the first record of the Japanese Tertiary foraminifers. The report of E. NAUMANN and M. NEUMAYI~ (1890) on the foraminifers from the Torinosu limestone of the Sakawa basin, Kochi Prefecture, is one of the few records of the Japanese foraminifers of the Jurassic Period.

With the beginning of the 20th century the results of Y ABE's researches on fossil foraminifers came to be published one after another, including the discovery of the Cretaceous Orbitolina limestone in Hokkaido (1901), the description of the Neogene orbitoids in Yamanashi Prefecture (1906), and the list of Late Cenozoic smaller foraminifers (1908); in the introduction of the last-named publication Y ABE stated, '' These minute fossils, though not so important geologically and paleontologically as other more highly organized types-for instance, Fusulina and its allies of the younger Palaeozoic, and Orbitoides and Nummulites of the older Tertiary-are too common to be quite ignored. As a full acquaintance of the detailed accounts of the Cainozoic stratigraphy which has received little attention, in the past, from our geologists, can be attained partly by a proper appreciation of the various factors of the fauna. I deem it by no means useless to

make a brief record of my observations on the minute forms of the Foraminifera." This introductory comment of Y ABE is interesting as it is indicative of the extent of the concern of scientists in those days about not only smaller foraminifers but also the Cenozoic stratigraphy. Y ABE's single-handed researches on foraminifers extended to fossils from Southeast Asia in addition to the Japanese specimens_ In the 1920's his successor Shoshiro HANZA w A began to take an active part in the foraminiferal researches, and a large number of papers were published under joint authorship of Y ABE and HANZA w A. The subject of their study was Order Foraminiferida although their energies were concentrated mainly on Neogene larger foraminifers. It was during this period that the first American journal on foraminiferal researches, Contribution from the Cushman Laboratory for Foraminiferal Research was published (1925). Yoshiaki OZAWA studied under J. A. CusHMAN, and their collaboration on Polymorphinidae bore fruit in the form of an excellent monograph that was published in 1930. CusH'viAN compiled the results of his study into the famous book Foraminifera: their Classification and Economic Use (1st ed., 1928). In the wake of this distinguished work, J. J. GALLOWAY published A Manual of Foraminifera in 1933. Thus, the academic world of foraminiferal research was given two great textbooks at one time.

Publication of joint papers by Y ABE and HANZA w A continued until the year 1935 when the Palaeontological Society of Japan was established. It happened to be the year when Kiyoshi ASANO started his active research on smaller foraminifers, and afterward the number of foraminiferal researchers increased. Taking these circumstances into accout, the writer would like to define the year


as the beginning of the second stage of micropaleontological study in Japan. It is true that the first stage had produced such records as Upper Paleozoic, Triassic and Jurassic radiolarians from the Sakawa basin of Kochi Prefecture by Singo YEHARA (1927), Plio:::ene ostracodes from the Kakegawa district of Shizuoka Prefecture by Jir6 MAKIY AMA (1931), and calcareous nannoplankton in the Shimajiri Group of Okinawa by HANZA w A (1925), but rese:uch work on these fossils made no progress before the second stage.

The second stage corresponds to the period from the time of the strained social situation facing the outbreak of the Pacific War to the dramatic termination of the War in 1945. This period was marked with the concentrative study of larger foraminifers by HANZA w A and the active research work of ASANO on smaller foraminifers. Their activities extended to the description, the stratigraphic distribution and the taxonomy of fossil faunas not only in japan but also over the vast Indo-Pacific region. One of the distinguished achievements of HANZA w A during this period was the study (1940) of larger foraminifers in the drill core from a deep well in Kita-Daitojima (North Borodino Island), and this is highly valued as a pioneer work of the post-war research on the structural development of the Pacific basin. ASANo's paper (1938) on fossil Nodosariidae is his representative work of this period. The number of researchers increased further during the second stage, and biostratigraphical survey of oil fields was pushed forward by Tsuneteru OINOMIKADO and others. Noticeable among these researchers was Kazuhiko IsHIZAKI who produced excellent papers (1939-48) on description and classification of smaller foraminifers while staying in

Taiwan. Unfortunately, however, he was destined to become the one and only japanese micropaleontologist who was directly victimized by the war.

Toward the end of the second stage, distribution and constituent species of diatomite deposits in Japan were summarized by Haruo OKUNO, and this paved the way for the post-war research on fossil distoms after the 50-year long gap since the work of BRUN and TEMPEim. The fossil radiolarians of the Sakawa basin that remained unstudied after the work of YEHARA were taken up by Toshio KIMURA, who described Upper Triassic-Lower Jurassic radiolarians (1944a, b, c). Teiichi KOBAYASHI and KIMURA (1944) published their general view on Paleozoic rocks and fossil radiolarian fauna of Japan. It was also during this period that B.L. CLARK and A.S. CAMPBELL were actively describing Upper Cretaceous, Eocene and Miocene radiolarians in California, revealing the stratigraphic merits of these fossils.

The transitional period between the second stage and the third stage experienced the worst publishing situation, and publication of micropaleontological papers reached the lowest ebb just as in other branches of science. In 1946, however, scientific journals began to be issued again. With the increasing opportunities for reporting the results in journals abroad the research activities became vigorous. In the case of foraminiferal research, it was after 1950 when the new series of the Transactions and Proceedings of the Palaeontological Society of Japan was started that the number of researchers increased and their activities became noticeable. In this period, ASANO summarized the results of his study on benthonic foraminifers and published "Illustrated Catalogue of japanese Tertiary Smaller Foraminifera" (1950-52).


The Catalogue gave a stimulus to younger researchers, and those who specialize in biostratigraphy gained in number. Consequently, the objects of study which had been limited mostly within the oilproducing regions along the coasts of the japan Sea came to include the entire Tertiary formations of Hokkaido, Honshu and Kyushu, and the results began to be published one after another since about 1960 (Yoshiro T AI, 1959 ; Manzo CHIJI, 1960; Shigeo MuRATA, 1961; Saburo IWASA, 1962; Torahiko INOMATA, 1962; Takashi MATSUNAGA, 1963; Yoshiki KIKUCHI, 1964; Yii HIGUCHI, 1964; Yasumochi MA TOBA, 1967; Naoaki AOKI, 1968). Many of these papers show a strong influence of KLEINPELL's " Miocene Stratigraphy of California" (1938) methodologically. AsANO also took the initiative in the study of the Cretaceous foraminifers in japan (ASANO, 1950a, b). Afterwards, the results of study of the Cretaceous foraminifers of Hokkaido were summarized into the monographic work (Yokichi TAKA Y ANAGI, 1960; Saburo YOSHIDA, 1963).

Study of Recent foraminifers was remarkably developed in the third stage. With the progress of researches on local fossil assemblages and the growing interest in biofacies and paleo-environment, the ecological investigation and study of dead assemblages and living assemblages in the Recent sediments were actively carried out, laying the foundation for clarifying the paleoecology. What had put spurs to these activities was the study by the group of scientists at the Scripps Institution of Oceanography. The group, with F.B PHLEGER and F.L. PARKER as the central figures, produced distinguished results in succession during a period from the 1940's to the 1950's. The group developed the short corer a:nd improved the vital staining method, and

these devices were adopted by the Japanese researchers and enabled them to make highly precise investigations. Prior to this, investigations of Recent foraminifers bad been made in Mutsu Bay (Yoshine HADA, 1931) and a few other bays and brackish lakes. In that third stage, however, the area of investigation expanded largely, and a large number of brackish, lakes, lagoons and bays, as well as the continental shelf and the upper parts of continental slope were studied by many researchers. Researches on biocoenoses are still few, among them are the valuable work of Yukio KuwANO (1962-63) on Pacific-side biocoenoses and that of MA TOBA (1970) on Matsushima Bay. Also, the achievements of Takayasu UCHIO (1960), who studied the living fauna off San Diego, California, are great.

With the progress in the ecological study, foraminiferal researchers came to take more interest in quantitative work in various fields, and mathematical analysis of fauna became popular. Hiroshi UJII:E and his students made faunal analyses of Recent and fossil benthonic and planktonic foraminiferal assemblages. Introduction of electronic computers made it possible to conduct high-order analyses, and distinguished results were gained (e. g., Tadamichi OBA, 1969).

In the latter half of the 1950's, H.M. BOLLI, W.H. BLOW and others set up fine biostratigraphic zonation in the Caribbean region by means of planktonic foraminifers, and this ignited the explosive increase of similar researches all over the world. The Miocene zonation by Tsunemasa SAITO (1963) was a comprehensive study in this field at that time, and on the basis of this zonation a precise correlation of Tertiary System in the low latitude regions was made for the first time. Afterwards, a large number of researchers have engaged themselves in


setting up zomtion of Paleogene to Pleistocene, and consequently the world-wide correlation is becoming more accurate. The biostratigraphic zonation, based mainly on marine sediments, has been further pushed forward, covering the vast region of western: North Pacific Ocean that includes the Pacific coasts of Southwest japan. Also, a scrutiny into the "global standard" of zonation is,under way. In the meantime, .t)le japan Sea coastal areas were taken up as the subject of biostratigraphic study, and the Late Cenozoic zonation, pec;uliar to japan located in middle latitudes, was proposed (Hisaya SHINBO and Seijuro MAIY A, 1971), presenting a topic relating to the history of development of the marginal seas in the Northwestern Pacific region. As for the Cretaceous stratigraphy, TAKAYANAGI (1965) attempted biostratigraphy of the Upper Cretaceous System of California on the Pacific coast. Lately, the biostratigraphic study using microfoJsi!s in deep-sea sediments is serving as a strong support to the development of the seafloor spreading theory and the plate tectonics. japanese researchers who participated in, or c;ooperated with, the JOIDES/DSDP project are publishing their results (e. g., Hiroshi UJIIE, 1975). They are expected to conduct active research work through Japan's participation in the JOIDES/IPOD project.

The results of researches on larger foraminifers that had been conducted ever since the first stage were compiled into the monograph on Micronesian foraminifers (HANZA w A, 1957), the voluminous book of "L.arger Foraminifera" (HANZA w A, 1968) and many other papers and books of HANZA w A, which formed a pyramid of this field. HANZA W A's SUC

cessors, though small in number, are concentrating their attention on the statistics of foraminiferal population and are

steady gaining results (Kuniteru MATSUMARU, 1971, 1973; UJIIE, 1966, 1973).

A series of studies by j. HOFKER, Sr. on the internal structures of foraminiferal tests greatly enlightened the Japanese researchers. Study in this field made a remarkable progress, irrespective of the types of foraminifers, that is, larger or sm9.ller and benthonic or planktonic (e. g., UJIIE, 1965-75). In the latter half of the 1960's, as the scanning electronic microscopy became popular, the study developed toward the elucidation of microstructures of test walls (TAKAYANAGI eta!., 1968). Moreover, measurement of paleo-sea water temperature by means of oxygen-isotope ratio (01" /018 )

was developed by H. C. UREY in 1947, which accelerated the study of paleooceans using planktonic foraminiferal tests by C. EMILIANI and others. OBA's work (1969) on the temperature changes of Pleistocene waters of the Indian Ocean contributed to the paleo-oceanography. Most recently, SAITO and j. VAN DaNK (1974) have proved that paleontologically inferred " planktonic " and " benthonic " foraminifers of Cretaceous and early Tertiary ages were actually planktonic and benthonic, on the basis of oxygen and carbon isotope measurements.

In the third stage, researches in the field of diatoms began with the study of biostratigraphic value of diatoms by W ataru ICHIKAwA, and elucidation of test structures, making the most use of electron microscopes, wa._s attempted (OKUNO, 1954-1959). Although 0KUNO did not get to reveal the phylogenetic significance of microstructures of diatom tests, his work is memorable in the respect that the electron microscopy was introduced to micropaleontology for the first time. Since then, the structural study of diatom tests has been continued .by W ataru IcHIKAwA, Kei OSHITE, Yukitoshi HAYASHI, Yasuo


HASEGAwA and other researchers, who recorded the fossil-coenoses in Upper Cenozoic fresh-water, brackish and rna. rine sediments of various parts of Japan, and reported the thanatocoenoses in Recent sediments. However, the methodological foundation of biostratigraphy was laid by Taro KANA Y A (1957) with his work on Californian Eocene diatoms. It was followed by his study of diatoms in the Miocene Onnagawa Formation (1959), which resulted in the establishment of biostratigraphic zonation of the Onnagawa Formation and corresponding strata. KANA Y A's work that gained worldwide frame was succeeded by ltaru KOIZUMI who set up the Upper Cenozoic zonation of the Oga Peninsula (1968). Despite of such progress, however, no start has yet been made with the study of pre-Miocene diatoms in Japan. On the other hand, the work of KANA Y A and KOIZUMI (1966) on the diatom thanatocoenoses of the North Pacific Ocean made a large contribution to the Recent diatom researches. This work clarified the distribution of diatom thanatocoenoses in Recent sediments, and it was on the basis of the distribution that KOIZUMI could accomplish his study of deep-sea biostratigraphy (Deep Sea Drilling Project) in the North Pacific Ocean (1973) and in the western marginal region of the Pacific Ocean (1975).

W ataru ICHIKAwA, while proceeding with his study of diatoms, recorded in 1950 the presence .of. silicoflagellates in the Miocene and Pliocene strata of Ishikawa Prefecture for the first time. Later, his collaborator A. BACHMANN (1964, 1967) made regular description and classification of silicoflagellates and archaeomonads in the Miocene diatomite of the Nato Peninsula. His work was followed by LING and others who described silicoflagellates and ebridians from the Miocene

Series of the Oga Peninsula (1971) and the Miura Peninsula (1972). As a result, the stratigraphic distribution of these groups of microfossils is becoming clear.

Radiolarian research in the third stage began with the work of Koichiro !CHIKA w A who made the first record of Paleogene radiolarians (1946) and successively described numerous species from the Permo-Triassic Sambosan Group (1951). Afterward, the study of Mesozoic fossils slackened until Akira Y AO (1972) started to work on spongosaturnalids. At present, the study in this field is expected to become more active. Biostratigraphic study of Miocene fossils was commenced from about 1954 by Kojiro NAKASEKO. Stratigraphic distribution of Neogene fossils in Hokkaido, in the region along the coasts of the Japan Sea, and in the joban and northern Kanto districts, was energetically pursued by NAKASEKO and his collaborators. On the basis of radiolarian fossils, they set up the zonation of the Neogene System along the japan Sea, mainly in the Toyama and Niigata basins, and revealed the faunal changes with geographic position and geologic time (1972, 1973). NAKASEKO (1964) also recorded radiolarian assemblages in the Recent sediments from the japan Trench. In the United States, on the other hand, study of fossils in the deep-sea sediments was started by W. RIEDEL and others at about the same time as NAKASEKO's work. The results of their stratigraphic study are being reported with the progress of the DSDP project. On the seas around the japanese Islands the Leg 31 report has been made lately by H. Y. LING (1975).

The presence of calcareous nannoplankton in the Neogene sediments of japan and in the Recent sediments of the neighboring waters was noticed during the first stage, but their description


cla::sifl ::ation and stratigraphic distribution remained unrecorded until the work of Toshiaki TAKAYAMA (1967) on the Late Cenozoic fossils in southern K1nto was published. From the next year, rese3.rches on nannoplankton were activated. Shiro NISHIDA an:l UCHIO p3.rticipated in the research work and many records were m1de on the Late Cenozoic and Recent sediments in various parts of Japan. The knowledge of the stratigraphic and geographic distributions of nannoplankton is rapidly gaining. In foreign countries, also, it was only after 1967 when the 1st Planktonic Conference (Geneva) was held, that the biostratigraphic study of this group of fossils made a remarkable progress. In 1971 E. MARTINI and D. BUKRY independently published the zonation of Cenozoic calcareous nannoplankton. Thus, the researches in this field came to boast of a great variety. Among the accomplishments of those days, the zonation of the youngest Cenozoic of Japan by TAKAYAMA (1973) and the Upper Cenozoic zonation of the Nansei Islands by NISHIDA (1973) are noteworthy. Published records of Paleogene and Mesozoic nannofossils are still few, but there are the reports on the Paleogene Setogawa Group (Susumu HONJO and Nachio MINOURA, 1968) and on the Upper Cretaceous Futaba Group (TAKAYAMA and lkuwo OBATA, 1968) ; and further developments of the study are expected. On the Recent species, a series of researches on the distribution of coccolithophorids in the Pacific Ocean were made by Hisatake OKADA and HONJO in the early part of the 1970's, which marked a milestone in this field of research. In the later half of the 1960's HONJO and his collaborators developed the electron microscopic technique for the study of microfossils. Their achievement ought to be rated high.

The first record of dinoflagellates and acritarchs was made by Kiyoshi TAKAHASH! in 1964 in his work on the fossils from the Oligocene Asagai Formation. It was in the 1970's that full-scale researches were carried forward, as exemplified by the reports on the Pleistocene, Plio::ene and Miocene fossils in Kyushu, Shikoku and Honshu by K. TAKAHASHI, Misaburo SHIMAKURA and others, and Kazumi MATSUOKA. Kenichi HARADA and his collaborators studied the fossils in the sediments of Black Sea (1973). A rapid progress in the biological and paleontological studies in the world was observed only after the time lapsed into the 1960's. Much expectations are laid on future studies in Japan, too.

Since 1866 when G.S. BRADY first recorded Japanese ostracodes, the ostracod researches in Japan focussed on living species and the reports were made mostly by biologists. Especially their work on bioluminescence attracted attention of the world scientists. As for the Cenozoic fossils, Tetsuro HANAI (1957-1961) in his serial works on ostracodes, entitled "Studies on the Ostracoda from Japan", dealt with living and fossil species comprehensively for the first time, and made efforts for their phylogenetic classification with proposal for several new subfamilies. This series includes "Historical review with bibliographic index of Japanese Ostracoda" (1959) which tells the status of researches in Japan up to that time. Description and classification of Late Cenozoic fossil assemblages were commenced in 1963 by Kunihiro ISHIZAKI. Synecologic researches on living bay fauna also were energetically carried out by him. Mesozoic fossils, however, have been little studied. In Japan, presence of some Cretaceous fossils is recognized, but no description has been given yet. The one and only achievement by Japanese


is the report on the non-marine fauna of the Sungari Group in Northeast China (HAN AI, 1951). The latest trend of the microp:lieontological re3earches in Japan is repre3ented by the fact that the study of microstructure of carapace arouses

considerable attention on its functional significance; moreover, the latest achievements at the ostracod study meetings of an international scale are spurring Japanese workers on to paleobiology.

Coelenterates

Makoto KA TO*

The Motsilji temple of northeastern Japan possesses a treasure called Jataiseki (serpent stone). This is a skeleton of colony of Permian tabulate coral known as Michelinia. It's weathered surface has a serpentine appeatance. This "treasure" is one of the instances of coelenterate fo3sils having attracted public attention since old times.

Scientific research of coelenterate fossils in Japan was started by M. NEUMA YR in 1890 when he described Convexastraea and Ch2etetopsis, Jurassic corals from the Torinosu Limestone. In 1902, Hisak::ttsu Y ABE described Permian Lonsdaleia from the Akasaka Limestone, and this was the first work by Japanese. Thereafter, study of fossil coelenterates was actively carried out by Y ABE and his students.

The history of study of fossil coelen-· terates in Japan can be divided into two periods, before and after World War II. The prewar period may be represented by sporadic descriptions of specimens. Though the number of researches was small, remarkable results were produced for various groups of coelenterates.

In the first place, Y ABE (1903) reported Stromatopora from Mesozoic beds, and

* Dc.Jartm~nt of G-eology and Mineralogy, Faculty of Science, Hokkaido University, Sapporo.

revised the previous idea which had restricted the occurrence of Stromatopora within the Paleozoic era. The research activities of Ichiro HAY ASAKA. Toshio SUGIYAMA and Kin'emon OZAKI, besides Y ABE, contributed towards revelation of fossil fauna of hydrozoans. mostly .stromatoporids, ranging from the Paleozoic System to the Cenozoic System of Japan, Korea and China. In particular, the work of OZAKI (1938) on the Chinese -Ordovician stromatoporids and the monograph by Y ABE and SUGIYAMA (1935) were distinguished achievements and largely influenced the coming researchers.

From 1915 to 1916 Y ABE and HAY ASAKA described Paleozoic coral fossils which were collected from various districts of Japan, China and Korea. The specimens examined by them included many new species and genera, and their reports served as important literature on coelenterates. A series of these papers contained no figure, but Atlas of Fossils in Geographical Research in China, 1911-1916 published in 1920 made up for the deficiency.

The research of the Omi Limestone by HAY AS AKA (1924) was the first attempt of biostratigraphic study of Japanese Paleozoic System by means of fossil coelenterates. The work was succeeded by



OzAwA (1925) who studied. the Akiyoshi Limestone. Later, Paleozoic corals from many parts of eastern Asia were described by Y ABE, SUGIYAMA, Motoki EGUCHI and others. Generally speaking, however, researches on fossil corals were less active than those on stromatoporids or other groups of coelenterates.

The oldest fossil-bearing bed in japan belongs to the Silurian System. From this bed Y oshio ONUKI detected Halysites, and SuGIYAMA (1940) wrote a monograph on corals and stromatoporids. Its discovery originated from OZAKI's study (1934) of Silurian corals of Korea. Nip. ponophyllum, a representative genus of japanese Silurian corals, was established then. With regard to tabulate corals, classification of Halysites by Y ABE (1915) is notable.

In connection with the classification of the so-called heterophyllids, Y ABE and SUGIYAMA (1940) proposed to divide Tetracoralla into Tetracoelia and Dicoelia. These subdivisions correspond to Order, so they are in the highest rank among taxa proposed by japanese for coelenterates.

Researches on Mesozoic corals of japan were conducted with Motoki EGUCHI as the central figure. The results were compiled by EGUCHI (1951) after the war, though the major parts were the products of researches made in the prewar days.

Cenozoic corals, including the living ones, are involved in the problems of paleoclimate, sea level fluctuation and reef formation, and so they have been the object of attention of geologists since old days. The controversy on the geologic age of the raised coral reef of Numa is especially famous. This reef is now assigned to Holocene, and is not regarded as a typical tropical or subtropical reef.

Y ABE and his students took interest

in the Recent coral reefs also, and carried out topographical, geological, biological and paleontological researches of coral reefs of the Palao Islands, collaborating with zoologists of the Palau Tropical Biological Station. These researches gave rise to many distinguished workers including TAYAMA, SUGIYAMA, EGUCHI and ASANO. The deep boring in Kitadaitojima conducted by YABE and others opened a new way of geological and paleontological research of coral reefs. Through these processes, the taxonomic study of Recent corals was advanced by paleontologists. The monographs on reefbuilding corals of japan and South Seas by Y ABE, SUGIYAMA and EGUCHI (1936, 1941) are the most remarkable products.

One of the interesting achievements is the work of MA (1934) on the seasonal growth of coral skeleton. His work was a herald of modern methods to be employed in the postwar researches such as the pursuit of the paleo-equator, the elucidation of continental drift, the chronological study, and so forth.

As an instance of a particular group, Conulariida was studied by HAY AS AKA and SUGIYAMA.

With the above-mentioned works at the peak, the first period of study came to an end. The war and the postwar chaos followed. Slackening of research activities was conspicuous.

The postwar period after 1945 began with the resumption and continuance of the prewar researches. In the postwar researches on fossil coelenterates, Masao MINATO took a leading role. His study of Paleozoic corals resulted in a monograph on Carboniferous-Permian corals of japan (MINATO, 1955), in which he defined a number of coralline fossil zones in the Upper Paleozoic System. His work on the ontogeny of Silurian corals of


Gotland Island of Sweden. was published in 1961. It introduced a new technique of study and at the same time exerted a large influence upon the later researchers. MINA To's wor\< and the Treatise on Invertebrate Paleontology, Part F, Coelenterata, whi!;:h was completed in 1956, served to bring up many researchers of Paleozoic corals. Those who engaged themselves in the study of fossil corals were; EGUCHI, Haruyoshi FUJIMOTO [ =HUZIMOTO], Hiroya GOTO, Takashi HAMADA, Wataru HASHIMOTO, HAYASAKA, Hisayoshi !GO, Ken-ichi IsHII, Tadao KAMEl, Kametoshi KANMERA, Makoto KA TO, Shigema KA w ADA, Michihiro KAwANO, Kenji KONISHI, Shiro MAEDA, MINATO, Masafumi MURATA, Takumi NAGAO, Mitsuo NODA, C. OKAMURA, Yoshio ONUKI, Masamichi OT A, OZAKI, C.L. ROWETT, Shigeo SAKAGUCHI, Toshihiko SA TO, Eitaro TAKAHASHI, H. TAKEDA, YABE, Y. YAMADA, Nobuo YAMAGIWA, Tsuruo YoKOYAMA and T. YOSHIDA.

Especially noticeable achievements were the serial studies on Halysitids by HAMADA (1956-59), the study of fine structure of corals by KA TO (1963), and the phylogenetic and paleogeographic rese:uches on Waagenophyllidae, Durhaminidae, Geyerophyllidae and Pseudopavonidae by MINATO and KATO (1965a, b; 1975a, b). New techniques of study, such as statistics and electron microscopy (SATO, 1963), were introduced.

Rezearches on Mesozoic corals became less active than in the first period, but Jurassic corals were studied by HASHIMOTO, MURATA, ONUKI, Kei MORI, and Triassic corals by KANMERA. Y AMAGIW A investigated the Triassic corals of Timor Island and South America. Thus, the researches on corals by japanese scientists covered the Pacific regions and the continents other than Antarctica and Africa.

In the study of Cenozoic and Recent corals, EGUCHI kept an . unchallenged position. HAMADA (1963) published an atlas of corals from Numa. Living coral specimens collected by the Emperor from Sagami Bay were compiled into a monograph by EGUCHT (1968).

OTA (1968) in his facies analysis of the Akiyoshi Limestone mentioned the structure of Paleozoic coral reef. MINATO and ROWETT (1968) discussed the way of reproduction of Paleozoic corals. Since 1968, KONISHI, Akio OMURA and others have been conducting dating of Quaternary corals by means of uranium isotope (KONISHI et al., 1975).

Study of Conularia began again lately by MURATA. On Mesozoic hydrozoans there are the research reports by HASHIMOTO. Stromatoporoids were neglected for a long time after the war, but MORI (1968, 1970) studied the Silurian specimens from Gotland Island of Sweden from the biostratigraphic and paleoecological standpoints, and completed a highly appreciated monograph.

Coelenterate fossils of japan leave much room for future study in view of description and taxonomy. With the introduction of various methods of study, interest of many researchers has overgrown the mere descriptions of fossils or fossil faunas and has turned towards the histology and functional morphology of skeleton, in other words a study of physiology and ecology of coelenterates, which would require the knowledge and technique of paleo-biochemistry, geochemistry and sedimentology.

The changes of hard tissue due to diagenesis and the process of fossilization are being pursued, as these will expand our knowledge of phylogeny of coelenterates and will also afford a clue to the natural resources of marine origin such as limestones and oil-bearing reef de-


posits. In 1971 the International Committee on

Fossil Corals and Coral Reefs was organized, and MINATO was elected as vicechairman. With this Committee in action,

exchange of information and cooperative researches became possible on a global scale. Future progress in this field must be remarkable.

History of Bryozoological Research

Sumio SAKAGAMI*

Study of fossil bryozoans commenced in 1924 when Ichiro HAY AS AKA reported five species of the Carboniferous bryozoans from the Omi Limestone. HAY ASAKA's work was succeeded by Yoshiaki OZAWA (1925) who described five bryozoans from the Carboniferous-Permian Akiyoshi Limestone. In either of the two papers, however, bryozoans were not the main subject of study as they were recorded only as accessory fossils in association with other important fossils such as fusulinaceans and corals. About fifteen years later, Toshio SUGIYAMA (1941) reported three species of "Batostomella" (specifically indeterminate) from the Hidaka mountains of Hokkaido and described them as Permian bryozoans. SUGIYAMA (1944) also described a Silurian bryozoan of Japan, Monotrypella? yabei (n. sp.), together with tabulate corals, etc. He was expected to take an active part in the study of the Paleozoic bryozoans, but unfortunately he passed away in 1944, the year his last work was published. Since then, nobody took any notice of the Paleozoic bryozoans in Japan until 1960, when Sumio SAKAGAMI (1960a, b) described two new bryozoan genera, Hayasakapora and NiP-

* Department of Geology, Faculty of Edu. cation, Ehime University, Matsuyama.

ponostenopom from the ]:lpanese Carboniferous and Permian systems, respectively. In the following year, descriptions and taxonomic researches of the Permian bryozoans from several localities of Japan were summarized by SAKAGAMI into a monograph. After that, he devoted his energies to the study of Carboniferous bryozoans, while concurrently supplementing his report on the Permian bryozoans with newly obtained data. Thus, he was able to clarify the Lower Carboniferous bryozoan fauna at Hikoroichi, Omi and Akiyoshi. At the 1st International Conference on Bryozoa held in Milan in 1968, SAKAGAMI read a paper entitled "Study on the Paleozoic Bryozoa of Japan and the Thailand-Malayan districts". In recent years, bryozoans from the Akiyoshi Limestone (Carboniferous-Permian) have been studied by Akihiro SUGIMURA (1974). Future studies are expected to reveal bryozoan faunas in the Silurian, Devonian and Upper Carboniferous systems of Japan.

The Mesozoic bryozoans in Japan have been studied very little. As for the Triassic bryozoans, only 30-odd species have been recorded in the world. Pseudobatostomella kobayashii was described by SAKAGAMI (1972) from the Carnic bed of the Sakawa basin in Shikoku. SAKAGAMI mentioned that one of the " Permian



Batostonzella " species reported by SUGIYAMA from Hidaka closely resembles Pseudobatostomella kobayashii, and he pointed out a possibility of the Hidaka specimen's being Triassic, not Permian. Occurrences of the Jurassic and Cretaceous bryozoans are known without any paleontologic studies as yet.

Study of the Cenozoic bryozoans in Japan was started by Katsuhiko SAKAKURA (1935, 1938) when he described Cheilostomata and Cyclostomata from the Pleistocene of Chiba Prefecture. After a gap of about twenty years, the Cenozoic bryozoans were recorded from the Pleistocene Daishaka Formation of Aomori Prefecture by J un KATAOKA (1957). Later, KATAOKA (1970)·described a large number of the Pleistocene bryozoans from the "Ryukyu Limestone" of Kikaijima, Kagoshima Prefecture. Recently, Tomoko HAY AMI is proceeding with the study of Japanese Cenozoic bryozoans, reporting their occurrences in the Pliocene of Okinawa Island and in the Mizunami Formation (Miocene) of Gifu Pre-

fecture, in addition to the northern part of Japan (HAY AMI, 1975).

The first paper on foreign bryozoans reported by Japanese was by Kin'emon OzAKI (1933), in which two species collected from the Toufangkou Limestone (Ordovician) of Province of Liaoning, South Manchuria were described. OzAKI's work was succeeded by Hisakatsu Y ABE and Toshio SUGIYAMA (1942) who described three species of the Permian Batostomella (Geinitzella) from Manchuria and Yunnan of South China.

Since 1962 the paleontological research in Southeast Asia is in continuance under the leadership of Teiichi KOBAYASHI. In some ten papers on the CarboniferousPermian bryozoans of Thailand and Malaya, SAKAGAMI has described more than 117 species of 27 genera. Thus, the relation between the Carboniferous-Permian bryozoan faunas of these regions and those of other regions is being clarified. An intermediate report of the results hitherto obtained has been made by SAKAGAMI (in TORIY AMA et al., 1975).

Brachiopodology in Japan-A Historical Review

Juichi YANAGIDA*

The first report on fossil brachiopods in Japan was the one by David BRAUNS (1881) which briefly described three species that were found, in association with molluscs, from the Quaternary beds of the Tokyo district.

A systematic study of fossil brachiopods began with the Permian specimens from the southern Kitakami mountain-

* Department of Geology, Faculty of Science, Kyushu University, Fukuoka.

land. After Hisakatsu Y ABE (1900) reported the occurrence of Leptodus under the name of Lyttonia, lchiro HAY AS AKA (1917, 1922, 1925) described 15 ·genera and 15 species including such peculiar-shaped ones as Leptodus and Richthofenia, and founded the basis of biostratigraphy of the Permian System of the Kitakami mountainland. Leptodus was reported also from the Neoschwagerina limestone of Kinshozan, Akasaka, central Japan, by HAY ASAKA (1925) who described 5



species in 3 genera including Scacchinella. Later, HA YASAKA (1932) added 3 species in 2 genera including Geyerella to the list of Permian brachiopods. Furthermore, 12 species in 10 genera, including such larger types as Orthotichia and Meekella, were described from the Nabeyama For. mation of the Kanto mountainland by HAY ASAKA (1933). From the Omi limestone HAYASAKA (1924) de3cribed 18 species in 8 genera of Carboniferous brachiopods, including Gigantoproductus and four other genera of Superfamily Productacea. On the basis of the result of this study he established fossil zones. HAY ASAKA's study not only confirmed the existence of the thick Carboniferous limestone but also contributed a great deal to the later biostratigraphical researches on the Carboniferous and Permian Systems. Owing to the efforts of HAY ASAKA and other pioneers, the knowledge of the Upper Paleozoic of Japan was widened and the yield of brachiopods increased rapidly. One of the noticeable achievements in those days was the discovery of the Devonian System containing Cyrtospirifer verneuili in the Kitakami mountainland by Y ABE and Mitsuo NODA (1933).

Researches on fusulinids contributed a great deal to gain the knowledge of the biostratigraphy of the Permian and the Carboniferous System. Brachiopods played an important role in clarifying the biostratigraphic successions of the Lower Carboniferous and the Devonian System of the Kitakami mountainland. Syringothyris and many other genera and species were identified by Masao MINA TO (1951, 1952, 1953) and Koichi T ACH!BANA (1956, 1962, 1969). Kitakamithyris of MINATO proved effectu1l in international correlation. Thus, the biostratigraphy of the Lower Carboniferous brachiopods of the Kitakami mountainland has ever since

held the position of the standard sequence in Japan. From the 1940's to the 1950's, taxonomical study of Silurian and Devonian brachiopods of Northeast Japan was conducted . by Toshiro SUGIYAMA (1942), Masahiro OKUBO (1956), HAY AS AKA and MINA TO (1954), and NODA and T ACH!BANA (1959); ·:.In and after the 1950's, Permian brachiopods of Northeast Japan were recorded in many papers including those by HAYASAKA, MINATO, Koji NAKAMURA, Jun'ichi T AZA WA and lchiro Y ANAGISA w A. The Lower Carboniferous brachiopods of the Kanto mountainland were described by Juichi YANAGIDA (1973).

On the Permian brachiopods from the Inner zone of Southwest Japan, excellent results were obtained by Sotoji IMAMURA, Daikichiro SHIMIZU, HAY ASAKA, Makoto KATO, and YANAGIDA. The Carboniferous brachiopods from the same region were recorded by YANAGIDA, MINATO, and KA TO, in their distinguished papers. In the Outer zone of Southwest Japan, Takashi HAMADA and NonA described Silurian brachiopods of Kyushu and Shikoku, and Permian brachiopods were described by YANAGIDA.

NAKAMURA (1972) described 25 species in 7 genera of Superfamily Davidsoniacea from the Permian system of the Kitakami mountainland, and presented a new idea about taxonomy and phylogeny of this superfamily. SHIMIZU (1961, 1963) clarified the period of occurrence of the Upper Permian brachiopod fauna in the Maizuru zone, and elucidated its habitat and the environmental changes with time, in connection with the coexisting faunas of other m::trine animals. YANAGIDA (1962, 1965, 1968, 1973) classified and described the Carboniferous brachiopods of the Akiyoshi area and made a detailed correlation. He also obtained a clue to the relationship between the sedimentary environment and the faunal changes with


time. HAMADA (1962) described 2 species of Lingula from the G2 stage (Wenlockian) and Conchidium from the Ga stage (Lower Ludlovian) of the Silurian System in the Outer zone of Kyushu; he maintained that the former suggests a calm environment and the latter an environment under agitated coral reef.

While the data of the Paleozoic brachiopods was thus accumulating, information on the Mesozoic ones remained surprisingly scanty. Edmund NAUMANN and Melchior NEUMA YR (1890) reported on the Mesozoic brachiopods collected from the neighborhood of the Sakawa basin, Shikoku. After that, there were reports by Singo YEHARA (1926), Y ABE and Saburo SHIMIZU (1927), Kango T A TEBA Y ASHI (1929), Teiichi KOBAYASHI (1931) and others, but many of them dealt with the materials from the Triassic and Jurassic Systems of Shikoku. Akira TOKUY AMA (1957-1958) examined the materials from the Triassic and Jurassic Systems of the Sakawa basin and surrounding areas and those from the Triassic System of the Chugoku region, and attempted a detailed correlation. He described new genera Sakawairhynchia and Spiriferinoides from the Triassic System and Naradanithyris and Neumayrithyris from the Jurassic System. He also expressed his view on grouping of the Triassic ribbed rhynchonellids.

On the Cenozoic brachiopods of Japan, there was a report by KOCHIBE (1882), which, in the form of an appendix to the above-mentioned BRAUN's report (1881), recorded 9 species in 6 genera of Tertiary brachiopods from the Kanto region. This work of KOCHIBE was, so to speak, the first of the academic papers on the Japanese Cenozoic brachiopods. Until the first half of the 1930's, brachiopodology in Japan was largely due to the achievements of HAY ASAKA (1932) and Matajiro

YoKOYAMA (1920). After the middle of the 1930's, Kotora HA TAI published the voluminous results of his researches on the Japanese Tertiary and Recent brachiopods. Especially his monograph, published in 1940, was a comprehensive record of his taxonomic study of the Tertiary brachiopods of Japan. HAT AI produced a l:lrge number of excellent results from his morphological and structural study of brachiopods. He (1941) also described fossil brachiopods from China and the Philippines.

In 1904 Y ABE wrote on the Devonian fossils from Hunan Province, China, and discussed the localities, horizons and affinities of brachiopod fossils among them. In the 1910's. Carboniferous and Permian brachiopods of China began to be reported by HAY AS AKA. This was the period when RICHTHOFEN's '"China" and MANSUY's monograph on the fossil brachiopods of Indochina and South China were published. HAY ASAKA (1922) described 41 species in 15 genera of Paleozoic brachiopods from Central and South China, and 11 species in 7 genera from the Lowest Permian System of North China. A marked affinity between fossil faunas of North China and brachiopods of the Ural-Timan region was pointed out.

From the latter half of the 1920's to the 1930's, Ryilji ENDO and KOBAYASHI continued their energetic study of paleontology and biostratigraphy of the Cambrian and the Ordovician of Manchuria (Northeast China) and Korea. A great number of species including many new ones were classified and described, among which Obolus and Lingulella were the major members. Their work was of a global achievement, greatly contributing to the establishment of the Lower Paleozoic biostratigraphy and the elucidation of the geologic history of East


Asia. The study was succeeded by Kinemon OZAKI (1931) who minutely classified and described the Lower Carboniferous brachiopods of Hunan province, China, and the Upper Carboniferous ones of Northeast China and Korea. The Devonian brachiopods of Northeast China were studied, though preliminarily, already in the early part of the 1940's by YABE, SUGIYAMA, KOBAYASHI and Juniti NONAKA. The materials collected by NONAKA from the Lesser Khingan region were lately re-examined in detail by HAMADA (1971). A precise correlation of brachiopods was attempted, and it revealed that they present a unique mixed fauna containing elements of other paleogeographic provinces.

Since the 1960's, paleontological survey parties have been dispatched abroad one after another, and the results of these efforts are accumulating with increasing speed. A series of paleontological researches in· Southeast Asia are particularly distinguished. The role played by fossil brachiopods in various parts of Southeast Asia is important. Notable works since 1964 are the following: HAMADA and KOBAYASHI made systematic descriptions of Lower Paleozoic

brachiopods from Thailand and Malaya ; YANAGIDA, HAMADA, Hisayoshi IGO and NAKAMURA did the same on Upper Paleozoic ones. Permian brachiopods from Cambodia were systematically described by NAKAMURA, KATO and Dong Ryong CHOI. These researches disclosed more than 120 species in about 80 genera which include such new genera as Echinocoeliop. sis (1968), Langkawia (1969), Malayanoplia (1969), Perakia (1969) and Swaicoelia (1968) by HAMADA, and Permundaria by NAKAMURA, KATO and CHOI (1970).

The history of brachiopodology in Japan is not long, being less than 100 years, but researches in some branches have risen to the world level. In the past, brachiopodology was mostly the means to clarify biostratigraphic successions, but nowadays the fossil brachiopods have begun to be studied from a purely paleobiological standpoint and the results are gradually budding out. For further progress, more specimens that were fossilized in their habitat and coexisted with various kinds of faunas must be gained. Such specimens might be found in reef limestones which are tLickly developed in the Upper Paleozoic.

Cephalopods

Ikuwo OBA T A*

The greater part of the history of researches on fossil cephalopods in Japan is occupied by the study of ammonites. A comprehensive history of the ammonites study in Japan has been published


by Tatsuro MATSUMOTO (1975) who divided the approximately 100-year long history into four stages (refer to Concluding Remarks by MATSUMOTO). It goes without saying that the history of researches in Japan owes a great deal to· the active works of Japanese scholars extending over several generations, but,



as MATSUMOTO points out, it is closely connected to the advancement of paleontology throughout the world. The history of cephalopod paleontology in Japan appears to reflect the characteristic development of paleontology in this country. It is expected that the Japanese cephalopod paleontology will be developed in a unique way by making the best use of the geographical and historical characters of the country, as well as by conducting international cooperation with a global field of vision and a sense of geobiology reinforced with modern techniques. Upbringing of cephalopod paleontologists (1873-1892): Early in the Meiji era (1873), a number of foreign geologists visited Japan and numerous fossils collected by them were sent to Europe for study by prominent scholars of those days. The results of the studies of these Japanese specimens were reported in scientific journals in various countries. One of the examples is the work of Edmund MOJSIsovrcs (1889) on Triassic cephalopods. In 1877 the Tokyo Imperial University was founded. Since then, Edmund NAUMANN and other foreign geologists served, in succession, as teachers at the university, bringing up Japanese geologists, conducting geological survey and making reports on fossils. Matajiro YOKOYAMA and Kotora JrMBO, who were educated during this period, went to Europe and studied under European geologists and paleontologists. Their reports on Japanese Cretaceous cephalopods were published in Germany (YOKOYAMA, 1890; JIMBO, 1894).

In foreign countries of this period, the classical monographs were completed already, as exemplified by the works of Alcide D'ORBIGNY (1840-42, 1842-51), Clement SCHLUTER (1871-76) and Ferdinand STOLICZKA (1863-65). It was also in this period that Albe~t OPPEL (1862) proposed

fossil zone, Alpheus HYATT (1868-1894) and W. BRANCO (1879, 1880) reported their studies of ontogeny and microstructure, Wilhelm W AAGEN (1869) announced the concept of variation with time, and K. A. VON ZITTEL (1876-93) published textbooks. Thus, at the start of cephalopod paleontology, Japan was already far in the rear of Europe for nearly 50 years. Cradle of cephalopod paleontology (1893-1915): YOKOYAMA received first lessons in paleontology from ZITTEL at the University of Miinchen. After returning home in 1893, he held a professor's chair at the Tokyo Imperial University and lectured on historical geology and paleontology. He published textbooks of paleontology in Japanese (1894, 1907, 1920), and also described fossil cephalopods (1904a, b), thus forming the basis of cephalopod paleontology in Japan.

Hisakatsu Y ABE, who graduated from the Tokyo Imperial University, studied in Europe. In 1912 at the newly established Tohoku Imperial University he commenced his studies of wide-ranging taxa from various parts of Asia. His study of cephalopods is represented by the reports that comprised the description of Pravitoceras (1901-02), of Nipponites (1903-04) and of Scaphites (1910). The results of researches during this period helped dating of Mesozoic formations. Between 1916 and 1920 there was a blank period in the Japanese cephalopod paleontology, probably due to direct and indirect influences of the First World War which lasted from 1914 to 1918. In Europe, however, some noteworthy achievements were recorded, such as the concept of biozone (R. WEDEKIND, 1916) and the presentation of paleobiogeographic province (Carl DIENER, 1916), in addition to descriptions and classifications of cephalopods.


Start of cephalopod paleontology (1920-1935) : This period was marked by the activities of Saburo SHIMIZU and a few other specialists who studied under Y ABE. Energetically they described Japanese cephalopods of different ages collected from various districts. At this stage, biostratigraphy based on cephalopods was attempted by Y ABE and SHIMIZU (1933) for the Triassic system, Seiichi MABUCHI (1933) for the Jurassic system, and Y ABE (1927) and SHIMIZU (1931, 1935) for the Cretaceous system.

In England of those days, the voluminous work of Leonard Frank SPATH, Gault Ammonoidea, was being published (1923-30, 1931-43), and Japanese researchers received the influence of SPATH more or less. SPATH carried out extensive researches on cephalopods of various ages, using the specimens collected not only in Europe but also in many other parts of the world. His classification is based on the concept of iterative evolution. It was in this period that DIENER (1925) published the Fossilium Catalogue of late Cretaceous ammonites, and 0. H. SCHINDEWOLF (1923, 1926, 1928, 1929, 1932-34) demonstrated his interest in ammonites. In Japan, on the other hand, the researchers came to take interest in the structure of cephalopods, and papers were written on siphuncle (SHIMIZU, 1929), on septa (Takumi NAGAO and Rinji SAITO, 1934) and on aptychus (NAGAO, 1931, 1932), besides descriptive, taxonomic and biostratigraphic papers. Fossil nautiloids of Cenozoic, Mesozoic and Paleozoic ages were also studied. Teiichi KOBAYASHI (1928, 1933, 1934, 1935) started the study of the Cambro-Ordovician nautiloid faunas from Northeast China and Korea introducing a new view on major classification as well as establishing genera. In America, A.F. FOERSTE (1921-35) published numerous papers

on the Paleozoic nautiloids. This period, therefore, may be called an epoch of diffusion of academic interest. Inactiveness and poor continuance (1936-1950) : This period, extending over the Second World War (1937-1945), produced no more than brief papers, owing to the forced interruption of research activities and the worsened situation of publication. Early Paleozoic nautiloids of China and Korea were continuously studied by KoBAYASHI (1936-38, 1940-42, 1947). Some Jurassic ammonites of Japan were also described by him (1947). Inside Japan, biostratigraphic efforts were continued, as represented by the accomplishments of MATSUMOTO (1942-43) and MATSUMOTO and Akira 0NO (1947).

In other countries during this period, various schools of learning were coming to the fore, such as the British sect represented by W.J. ARKELL (1933, 1935-50), the French sect represented by Eliane BASSE (1947) and Maurice BREISTROFFER (1940, 1947) under Charles JACOB, and Maurice COLLIGNON (1948-50), and the America! sect represested by R.W. IMLAY (1939, 1942, 1943, 1948), A. K. MILLER and W.M. FURNISH (1937, 1938, 1940), etc. Paleozoic nautiloids were energetically studied by Curt TEICHERT (1939, 1940), R.H. FLOWER (1936, 1938-1943, 1945-1950), etc. The level of cephalopod paleontology in Japan at that time fell much behind the world level. Activation and internationalization (1951-1958) : In this period the international exchange of knowledge was reopened and became active, supported by the young researchers studying abroad and the mature scientists visiting overseas countries, as well as by the improved situation for publishing academic journals. The volume on ammonites in Treatise on Invertebrate Paleontology was published in 1957. At the same time, de-


scription, classification and biostratigraphic division of Japanese cephalopods developed remarkably. A great many papers were written on Paleozoic, Mesozoic and Cenozoic cephalopods, in an effort to catch up with the level of the western countries. Especially notable of this period are that the publication of the monograph on the Cretaceous ammonites by MATSUMOTO (1954-57) kept on, that the description of the Jurassic ammonites by Tadashi SA TO (1954-58) progressed, that the Cretaceous belemnites were studied by Tetsuro HANAI (1953), and that the Japanese Tertiary nautiloids were described by KOBAYASHI (1954-58). KOBAYASHI is highly commendable for his longstanding studies of various kinds of nautiloids from Asia. In foreign countries, leading figures in the academic world were making their appearance here and there, namely, C.W. WRIGHT (1952, 1953, 1955, 1957) of London, R.A. REYMENT (1955, 1956, 1958) of Stockholm, J. SORNAY (1955) of Paris, SCHINDEWOLF (1951, 1953, 1954, 1957, 1958) of Ti.ibingen, Bernhard KuMMEL (1954-57) of Harvard, V.N. SHIMANSKIY (1954-57) and Z.G. BALASHOV (1956, 1957) of U.S.S.R. Rapid progress and new paleontology (1959-1973): The characteristics of cephalopod paleontology in this period may be represented by the following three facts, not to speak of the marked increase in the number of descriptive and taxonomic papers. The first fact is that the biostratigraphic division of the Japanese Mesozoic was established at the international level, in consequence of accumulation of materials. For example, the division of the Cretaceous system by MATSUMOTO (1959, 1969), of the jurassic system by SA TO (1962) and of the Triassic system by Yuji BANDO (1964, 1967) was published. Especially the fossil zo-

nation by MATSUMOTO was quite unique, the first of its kind in the world. He established the zonation by consciously discriminating three types of ammonites, namely, the fiat and smooth thin-shelled type that predominates in the relatively pelagic, muddy, geosynclinal facies of the circum-Pacific region, the ornamented thick-shelled ammonite predominant in the neritic facies of the continental shelf region, and the type represented by abnormally coiled Baculites and Scaphites that predominate in the facies intermediate between the above two. Thus, his zonation is going to play an important role in linking the circum-Pacific region to the Tethys Sea for a global correlation. The second fact is that the research work of the japanese paleontologists came to deal with ammonite assemblages and type specimens from foreign countries, in parallel with the popularization of air transportation and the economic development of Japan. The resultant numerous papers contributed to the international correlation and the knowledge of paleogeography of the world. A typical example is the work of MATSUMOTO (1959-60) on the Cretaceous ammonites of California, which was rated high abroad as indicative of the high level of the japanese researchers. The third fact is that the biological study of fossil cephalopods was commenced. For example, Ikuwo 0BATA (1959, 1960, 1965) applied the relative growth formula to ammonites for the first time, and distinguished minute changes in the growth pattern. This work paved the way for the study of relative growth of ammonites by the researchers in the western countries. MATSUMOTO (1963) related the difference in habitat by the ammonites genera to the pattern of ontogeny and phylogeny. Takayo Funw ARA (1961) studied amino acids in am-.

A Concise Histo'ry of Palaeontology in Japan 37

monites. The volume on nautiloids in Treatise

on Invertebrate Paleontology was published in 1964. In foreign countries, original ideas and techniques were introduced one after another, such as the simulation model for examining morphology and function of fossil cephalopods (David M. RAUP, 1965-67), the combination of relative growth and statistic technique (S.J. GouLD, 1966), the experiments on buoyancy of living cephalopod shell (E. J. DENTON and J. B. GILPINBROWN, 1961, 1963, 1956, 1973), the paleobiochemical study (P. H. ABELSON, 1954, 1955, 1956, 1963), the functional morphology of cephalopods and statistics (REYMENT, 1956, 1973), and the study of cephalopod shell by means of a scanning electron microscope (H. K. ERBEN et at., 1969). Efforts for the taxonomic description and the systematics were continued (e. g., William A. COBBAN, 1961, 1962, 1964, 1965, 1969, 1970, 1972; J.A. JELETZKY, 1966, 1969; Raymond CASEY, 1961-66; Jost WIEDMANN, 1960, 1962, 1963, 1968-70; COLLIGNON, 1959-73; KOBAy ASH!, 1959-61, 1971). Nevertheless, the scientific interchange between the foreign countries and Japan was not in a satisfactory state. The rise of paleobiology (from 1974 on): The characteristics of this period are essentially the same as, and continuous from, the preceding period. Paleobiological study of fossil cephalopods in Japan was given a stimulus and the level of young researchers rose, bearing comparison with the western countries. The rise of paleobiology in Japan owes a great deal to Itaru HAYAMI (1969-71) with his efforts in the statistical treat-

ment of relative growth of fossil molluscs, as well as in the study of their population genetics and life range. Statistical technique was adopted also by Hiromichi HIRANO (1975) in his study of the shell structure of Cretaceous Gaudryceras in its early stage of ontogeny and its growth pattern, and by Kazunari T ANABE (1975) who studied the relative growth and functional morphology of Cretaceous Scaphites. Standing on the sound basis of biostratigraphy, these people are continuing their research work, enduring the criticism from RAUP, REYMENT, GOULD and others. Future progress in this field is expected. International cooperative study from the paleobiological standpoint has been started. Reiner JoRDAN and MATSUMOTO are working on pathological phenomena of ammonites, ERBEN and OBATA are studying cephalopod shell structure by means of scanning electron microscopy. REYMENT (1974) analyzed a generic level transition from Subprionocyclus to Reesidites mainly on the Japanese material.

In 1976, an international meeting and a forum are to be held in Hokkaido, with the object of discussing " MidCretaceous Events" which was adopted in 1970 as part of the International Geological Correlation Projects. The present level of cephalopod paleontology in Japan, especially with regard to description, classification, biostratigraphy and paleobiology of late Cretaceous ammonites, is as high as the level in the western countries. Researches on fossil cephalopods in Japan will be further internationalized and will keep on advancing.


Fossil Molluscs

Tsugio SHUTO*

Paleontological study of Japanese molluscan fossils, excluding cephalopods, began with the work of D. BRAUNS (1881) who dealt with Quaternary fossils from southern Kanto. It was followed by the reports of E. NAUMANN and M. NEUMAYR (1890) and Matajiro YOKOYAMA (1890, 1910, 1911). But it was YOKOYAMA (1920) who initiated a systematic study of fossil molluscs.

In those days researches and papers were centered upon Cenozoic and Mesozoic molluscs, whereas information on Paleozoic molluscs was extremely scanty because specimens were limited in both quantity and quality. In addition to this unbalance of available information, there was a tendency among the researchers to set their goal at elucidation of biostratigraphy, and so the research work was carried on by separate groups for a specified geologic age. Consequently, little attempt was made to put together all available information and material obtained by those specialized in Cenozoic, Mesozoic or Paleozoic molluscs, so as to compose a phyletic evolution or to establish a system of classification. However, after the war, especially in the 1950's, an idea that fossils should be regarded as "organisms" began to spread among the researchers while proceeding with their study in the respective fields. Based on this idea, the results of researches in different geologic ages came to be linked up gradually, and the linkage is apparently growing ever since the


1960's. Therefore, the history of fossil molluscan researches in Japan may be divided into two periods, from 1881 to 1952 and from 1952 on, although the boundary is not sharp as the two periods partly overlap, and it may be admissible to define the first period as a period of biostratigraphic study and the second period as that of biological study. In view of the participants of the respective works, it can be said that the first period was marked with research activities at Tokyo, Tohoku and Kyoto Universities, and the second period is the time when the center of study has been aplit into multiple institutions. It goes without saying that there were some excellent pioneer works in biology during the period of biostratigraphic study, and biostratigraphic researches are continued, with higher precision, in the present period of biostratigraphic study. This article reviews the history according to the respective series of researches.

Series of biostratigraphic study: Academic study is apt to develop through tense competitions among researchers or research groups. This applies to the fossil molluscan researches in Japan. The above-mentioned three universities that took the lead in biostratigraphic study, particularly during the former half of the first period, seem to have worked with different subjects and methods of study. That is, at Tokyo University description and biostratigraphy of Cenozoic molluscs were dealt with, and biostratigraphy of Mesozoic and Paleozoic molluscs at Tohoku University, whereas Kyoto Uni-



versity had an inclination toward more precise biostratigraphical and biological researches supported by detailed field work.

M. YOKOYAMA published his voluminous works on molluscan fossils from the Miura Peninsula and its vicinity (1920) and those from Shimoosa and Kazusa (1922). After that, he came to deal with late Cenozoic molluscs. The area of his study was vast, extending from Sakhalin to Taiwan, and the subject material ranged in age from younger Quaternary to Paleogene, but his efforts were concentrated on the study of the latter half of the Cenozoic era. His detailed reports on the fossil molluscs of southern Kanto are distinguished accomplishments. The results of his energetic descriptions disclosed the outline of late Cenozoic molluscs of Japan and laid the foundation of the present-day researches. A little later than YOKOYAMA, Jiro MAKIY AMA started his study of younger Cenozoic molluscs, and he employed a new method worthy of note. He restricted the area of his study to a certain district where he carried out precise field survey and collected specimens, particularly in the Kakegawa district. On the basis of the material thus obtained, he endeavored to establish detailed biostratigraphy (1925, 1927, 1931, et seq.). His reports on the Miocene molluscs from the Korean Peninsula (1926, 1936) deserve special mention. He also introduced the biological concept and technique into the study of fossils, as will be described in the later section. Takumi NAGAO studied at first the Cretaceous molluscs under the guidance of Hisakatsu Y ABE, but he later extended his research work to the Paleogene molluscs. His comprehensive description (1928, et seq.) of molluscs from coal fields in Kyushu on which the Paleogene biostratigraphy was established proved an

important contribution. When the research activity of YOKOYAMA was about to cease, Shichihei NoMURA and Kotara HAT AI began to study molluscs of younger Cenozoic age and energetically described the major fossil faunas of the Tohoku district and many other parts of Japan. NOMURA published the results of his laborious work on the fossil molluscs from Taiwan (1933, 1936) and on fossil and living Pyramidellidae from Nansei Shoto. HAT AI's description of living molluscs from Micronesia was also a distinguished achievement. Yanosuke OTSUKA, like MAKIYAMA, endeavored to establish detailed field stratigraphy and biostratigraphy, and produced many noteworthy results as represented by the study on the northern Kitakami massif (1934) and the Ochi Graben (1935). Around 1940 Koichi SUZUKI described Paleogene fresh-water molluscs from Hokkaido and Kyushu, and Ken'ichiro OT A TUME described Paleogene freshwater molluscs mostly from Hokkaido. In comparison with the 1930's, the number of papers decreased during the wartime days, but it was characteristic that some papers dealt with the specimens obtained outside the country, such as those from the South Sea area (Ichiro HAY ASAKA, et seq.) and from the Asian Continent (OTSUKA, 1942; SUZUKI, 1942).

The study of Cenozoic fossil molluscs was greatly pushed forward by the results of the researchers of living molluscs. Publication of J. THIELE's Handbuch der systematischen Weichtierkunde, Bd. 1 (1931) and Bd. 2 (1935) made an epoch in the taxonomic study of molluscs of the world, and with this as a turningpoint the modern phylogenetic classification began to spread in Japan. The works of W. WENZ (1938-44) and A. ZILCH (1959-60) in the form of Gastropoda (Handbuch der Palaeozoologie, Bd. 6)

40 "MATSUMOTO, T. et al., edit.:

afforded an easier approach to the taxonomic study of fossil gastropods which the researchers had been liable to shun. Tokubei KURODA is known as a leading figure among the japanese re3earchers on living molluscs, and his results largely promoted the phylogenetic classification of molluscs in Japan, though qis description of fossil molluscs from Shinano should not be overlooked either. In this field, contributions of Kotora HA TAI, Tadashige HABE and Katsura OYAMA are also remarkable. The works that played an important role in spreading the Cenozoic molluscan researchers after the war are ; the reclassification of fossil molluscs from southern Kanto (Iwao TAKI and OYAMA, 1954), the reclassification of Cenozoic molluscs from southern Kanto (MAKIYAMA, 1957-60), and the re-examination of the japanese Paleogene molluscs (OYAMA, Atsushi MIZUNO and Toru SAKAMOTO, 1960).

The termination of the war was the return of spring to the molluscs researchers too. Differing from the prewar days, the center of research was not restricted to the afore-said three universities, and a great many researchers at numerous institutes competed with one another in producing excellent results. However, most of research works were intended for biostratigraphy of different districts, and were hardly expected to give rise to phylogenetic classification, evolution and paleoecology. The study of the Cenozoic molluscs that had been apparently prosperous for some time slowed down rapidly within 10-odd years after the war when the local biostratigraphy was settled to some extent, and many researchers parted from molluscs. Around the year 1960, stratigraphic and descriptive studies of the japanese Cenozoic molluscs were nearing the saturation point, as it were, with accumulating re-

cords of description and re-examination of numberless local assemblages of fossil molluscs. Representative works in this field were; Kiyotaka CHINZEI on Sannohe (1959, 1961), Shozo HAYASAKA on the Atsumi Peninsula (1961), Katsumi HIRAYAMA on Arakawa (1954) and Asagai (1955), Junji ITOIGA WA on Mizunami (1955-63), Takehiko Iw AI on Tsugaru (1964), Yasuhide IWASAKI on Shiobara and corresponding areas (1970), Yasuhiko KAMATA on joban (1962), Saburo KANNO on Chichibu (1960), Yoshio KASE;\10 and Nobuomi MATSUURA and Onma (1965), Koichiro MASUDA on Toinnai (1966), Hiroshi OZAKI on Choshi (1958), Tsugio SHUTO on Miyazaki (1955-62), Kunio TAN AKA on Shinano (1959-60), Karyu TsuDA on the Kurosedani formation (1959-60), Satoru UOZUMI on Hokkaido (1957-66), and MACNEIL on Okinawa (1960). In the meantime, fossil molluscs of foreign countries, mostly of Southeast Asia, were began to be described, as exemplified by the works of KANNO (1973-75) and KoTAKA (1975) with the specimens from Taiwan, and of IwASAKI (1970) and SHUTO (1969, 1971) with the Philippine specimens. Specimens from Yakataga, Alaska were studied by KANNO (1971), and Pectinids of North America and North Pacific Ocean by MASUDA (1970-73). These studies, however, have not yet reached the stage to establish satisfactory biostratigraphy and paleogeography.

Phylogenetic classification is essentially a comprehensive task, and so it ought to be mentioned in the section of biostratigraphic study. But it is touched upon here because the proposed classification is closely related to stratigraphic paleontology. On account of the actively accumulated data, phylogenetic examination of taxonomic groups of various sizes became possible in the 1960's. There were, of course, some forerunning studieo


in this field, such as the works of Takumi NAGAO and Kazuo FUJIOKA (1941) on Acila and of Norio lKEBE (1942) on Calliostoma. Notable among the postwar achievements were the results of Kazuyoshi IDA (1952) and Tamio KOT AKA (1959) on Turritella, UOZUMI (1959) on Yoldia and Portlandia, Koichiro MASUDA (1962) on Pectinidae, Koji NODA (1966) on Anadara, and Tokio SHIKAMA (1969) on Fulgoraria.

The check list of the Japanese Cenozoic molluscs (HATAI and Shozo NISHIYAMA, 1952) and that of the Japanese living molluscs (Tokubei KURODA and Tadashige HABE, 1952), published soon after the end of the war, proved very useful and greatly facilitated taxonomical and biogeographic researches of Cenozoic molluscs. Perhaps these check lists marked a suitable end of the period of biostratigraphic study.

Study of Mesozoic molluscs was started by YoKOYAMA (1890) with his stratigraphic descriptions of Cretaceous specimens from U rakawa and other parts of Hokkaido, but it did not become systematic until Shingo Y. EHARA reported on Trigoniinae (1915, 1923). Since Y. EHARA's work, pelecypods of Palaeoheterodonta, such as " Trigonia " and "Unio ", became the major subjects of study of Mesozoic molluscs by a large number of researchers, so that it appeared as if these pelecypods represented Mesozoic molluscs. The prewar and wartime achievements along this line, besides the above-mentioned Y. EHARA's work, were the reports of Teiichi KOBAYASHI and Koichi SUZUKI on Rakuto [Nagdong]-Wakino (1936), Tetori (1937) and Yamaguchi (1939), of Y ABE and Zi-iti HAYASI on Manchuria (1938), and of SuZUKI on Shiragi [Shiira] (1940), on the Sungari series (1941) and the summary of East Asian molluscs (1949); SuzuKI's

contributions were especially notable. After the war, researches in this field were expanded by Shiro MAEDA (1949, et seq.), Mitsuo NAKANO (1957, et seq.), Minoru TAMURA (1959, et seq.), Yoshihisa 0HTA (1959, et seq.), ltaru HAYAMI (1962, et seq.), and S.-Y. Y AN (1974). OHTA also re-examined corbiculids of North America.

As for pelecypods of Pteriomorphia, their stratigraphic value had been recognized since early days, and much efforts were made in the study of them. The Cretaceous " Inoceramus" among them was cl:l.ssified and stratigraphically studied by Takumi NAGAO and Tatsuro MATSUMOTO (1939-1940), and on the Jurassic specimens similar results were obtained by I. HA YAMI (1960). The Triassic Pteriomorphia was first taken up by T. KOBAYASHI (1935), and the study was largely developed after the war by Koichiro lCHIKA w A, TAMURA, and Akira TOKUY AMA. The fact that the researches on Mesozoic molluscs were concentrated on the above-mentioned taxonomic groups may be partly ascribed to their abundant occurrences, but their valuableness as index fossils must have been the major reason.

Collective description of fossil assemblages, treating them as a molluscan fauna so to speak, commenced with the work of Y ABE and NAGAO (1925) who studied chiefly Cretaceous marine molluscs. Faunal description of molluscs advanced rapidly after the war, giving rise to the study of paleogeography. The Triassic molluscs were studied in succession by NAKAZAWA, ICHIKAWA, TAMURA, TOKUY AMA, the Jurassic ones by TAMURA, HAYAMI, MAEDA, and the Cretaceous ones by HAY AMI and ICHIKAwA and Yasuo MAEDA. Especially, the systematic study by TAMURA and HAY AMI is rated high.


Mesozoic fresh-water molluscs of foreign countries had been studied by japanese paleontologists in the prewar and wartime days, as mentioned before. After the war many papers on marine molluscs were written by TAMURA (1968, et seq.), HAY AMI (1968, et seq.), and others in Geology and Palaeontology of Southeast Asia. In the Triassic system of Thailand and Malaya, fossil faunas are being recognized (KOBAYASHI and TAMURA, 1975).

Ichiro HAY ASAKA was the first to describe Paleozoic molluscs of japan in his study of pelecypods from Kitakami and Ogachi (1923). Study of Paleozoic molluscs made a start a little later than that of Mesozoic and Cenozoic ones, but its progress was extremely slow. In 1925 HAY ASAKA described part of the molluscs from Akasaka Kinshozan, and published a sequel of it in 1943. HAY ASAKA's work was the one and only example of the systematic description of Japanese Paleozoic molluscs during the prewar to wartime period. In 20 postwar years the molluscs from Kitakami were described by Masabumi MURATA (1964) and the molluscs in the Akiyoshi limestone were described by SHIKAMA and Tamio NISHIDA (1968) and NISHIDA (1968). Under these circumstances, the work of NAKAZAWA and N.D. NEWELL (1968) in summing up the Permian molluscs of japan was of great value. Molluscs of foreign countries were mentioned by KOBAYASHI (1930, 1931) in a part of his description of Ordovician faunas of Korea and Manchuria. There were some other brief descriptions of them.

Series of biological study: Commencement of biological study of fossil molluscs dates back to the prewar days. Using fossil and living Umbonium, MAKIY AMA (1925) attempted to establish the evolutional system, and inferred the

evolutional relations among the genera of Umboniinae from the morphological development of their shells. Later, he discussed the series of evolution of Siphonalia with the similar method (1941). MAKIYAMA (1900) also tried to analyze variation and affinity of populations by means of statistical treatment of Glycymeris yessoensis from different localities. His work was by far advanced at that time, and was followed by not a few researchers in pursuing the evolutional series of Umbonium with additional material. Nevertheless, the importance of morphological development as a phase of ontogeny was not fully understood by researchers in general, and morphology was seldom put to practical use in classifying and describing molluscs. It was only after the war that the significance of statistical and biological researches was duly recognized and put in practice. The same situation applies to the significance of pelecypod fossils perforated by predatory gastropods, as pointed out by I. HAY ASAKA (1933). With these forerunning works in the background, the results of biological researches began to be published after the war, particularly from about 1953.

The year 1959 happened to be the lOOth year since the publication of The Origin of Species by Charles DARWIN. This gave an impetus to enlightenment movements and panel discussions on evolution. One of the consequences of these activities during the 1950's was the publication of " Seibutsu Shinka " (Evolution of Life), its first issue appeared in October 1953. The journal offered a common ground to biologists and paleontologists for joint discussion. The first wave of this stream was represented by the works of KaT AKA on Anadara granosa (1953) and Turritella (1954) and Masae OMORI and Tsutomu UTASHIRO


on Pecten albicans (1954), who attempted to pursue population changes and evolution through statistical treatment of living and fossil molluscs, and the work of ICHIKAWA on Entomonotis (1954) in pursuit of morphological variation of groups with time. It cannot be denied that the results of biological studies which were still in the budding stage are poorer than the contemporary results of classification and description having the long-standing tradition, but the abovementioned works are worthy of notice as being indicative of a trend of future development. In the course of these studies, the importance of speciation and its mechanism as a vital point of evolution came to be recognized, and attempts were made to have a correct understanding of speciation by means of genetic and mathematic analyses of population. I. HAY AMI improved the mathematic method (1969, 1970, with Akihiko MATSUKUMA) and paved a way for paleogenetics (1973), thus providing a large possibility for future studies. The excellent analysis of Inoceramus (Sphenoceramus) naumanni by Kazushige T ANABE (1973) stands on the same basis as above. In dealing with the mechanism of speciation, there is developing a trend to grasp speciation ecologically, in view of such biological concepts as sympatric and allopatric populations (T. SHUTO, 1957), although any satisfactory results are yet to come. SHUTO's work (1974) discussing the relationship of embryonic ecology and spaciation of gastropods may be placed along this line.

In parallel with increasing efforts in the study of evolution, researchers' interest in paleoecology was growing. It must be noted that the Matsukawaura Research Group, a leading figure in the early part of the history of paleoecological research, pointed out the importance of compara-

tive study of living and fossil organisms, and the consequent significance of taphonomy (1954). Although no succeeding works are reported as yet, taphonomy is one of the challenging problems for future studies. Paleosynecology, as a branch of paleoecology, had been studied since early days, but many of the papers showed an inclination toward environmental discussion (ITOIGAWA, 1959; TsuDA, 1960; SHUTO, 1960; MASUDA, 1962; etc.). It may be a natural consequence that these studies were developed in the manner of integrating paleogeography and biostratigraphy, rather than aiming at Paleosynecology itself. Studies in this field are represented by the works of CHINZEI and IWASAKI (1964), who analyzed differentiation and variation of the Neogene molluscan faunas of the Tohoku district, and of Shigeru AOKI (1960) and CHINZEI (1963) laying stress on variation of assemblages. Faunal analysis from association of species, feeding habit and life form was attempted by SHUTO and Shigemi SHIRAISHI (1971), which made one step toward functional analysis of ecological community, but it leaves room for doubt that to what extent the analysis of molluscs alone can be effective. An approach to paleosynecology was shown also by KoTAKA (1973) who employed the statistic technique for analyzing intraspecific assoc1at10n. The works of HABE (1956) on thanatocoenosis in bays and of Qy AMA (1950, 1953-1954) who compared living and fossil communities served as a flywheel for paleosynecologic study.

Paleoautoecology has been studied on the basis of individual life form or behavior, feeding habit and functional morphology. Life form or behavior of pelecypods was studied by ITOIGAWA (1963) on their wood-boring, by MASUDA (1968-1972) on rock-boring, by UOZUMI


(1956) on borrowing and boring, and by Iwao KOBAYASHI and OMORI (1973) on pelecypod shells bored by coexisting polychaetes. On feeding habit, Makiko KOBA Y ASH! and l. KOBA Y ASH! (1971) examined the relation between the predator and its boring act, I. HAY AMI (1969) studied the functional form of Mesozoic planktonic pelecypods, and TANABE (1973) discussed the life form of Inoceramus from the viewpoints of its shape, shell structure and relative growth.

Histological study and biochemical study of fossils were introduced roughly concurrently into the field of molluscs paleontology. In the 1960's microstructures of pelecypods were studied by OMORI, I. KOBAYASHI and Matsutaro SHIBATA (1962) and their biochemistry by Masahiko AKIYAMA (1964). From the start the microstructures were discussed in connection with functions (OMORI and KOBA Y ASH!, 1963 ; SHIBATA, KOBAYASHI and Hisashi KAIBARA, 1968; OMORI, 1971). Study of the microstructures was also

united with biochemistry of organic matter, and is being proceeded, heading towards clarification of the shell-forming mechanism (UOZUMI and Keiji IWATA, 1969 ; AKIYAMA, 1971 ; Hiroyuki HAT ANO, 1971). Also, elucidation of diagenetic changes of conchiolin is under way (IWATA, 1975).

When the 180 method was employed for paleoclimatological analysis to obtain information on paleotemperature of waters, pelecypods were utilized as part of the working material (Hitoshi SAKAI, Kenji KONISHI and Osamu NAKAMICHI, 1969; Sumio HORIBE, Nobuaki NIITSUMA and Toyosaburo SAKAI, 1969; Fujio MASUDA and Kazuhiro TAIRA, 1974).

Biological study of fossil molluscs made a remarkable progress in spite of its young history. For some groups of molluscs, however, researchers are still fumbling in the darkness. so to speak. Really useful biological results are yet to be produced.

Arthropoda

Takashi HAMADA*

Trilobita: Trilobite researches in japan can be divided into two groups, study of japanese specimens and study of specimens from abroad. When the history of japanese trilobite researches is reviewed by splitting it into four periods, the first period is regarded as the dawning period. The first record of trilobite is found in "Die japanischen Inseln" by Toyokichi

* Department of Earth Sciences and Astronomy, College of General Education, University of Tokyo, Tokyo.

HARADA (1890), which mentioned PhilliPsia-ahnlichen Trilobiten from Obama of the Kitakami district. Afterwards, Permian trilobites were reported by Kotora ]IMBO (1896, 1902), lchiro HAY ASAKA (1933), and Takumi NAGAO (1931).

The second period was, so to speak, a period of development, represented by the works of Hisakatsu Y ABE and Toshio SUGIYAMA (1938, 1940, 1941, 1944) following the discovery of Siluro-Devonian system in the Kitakami mountainland. Phacops, Encrinurus and other trilobites



were reported from a stratigraphic standpoint. Carboniferous Palaeophyllipsia SUGIYAMA and OKANO (1944) was the first trilobite genus established on the basis of japanese material.

The third period, from 1950 to 1972, is marked by accumulation of data. As the occurrence of the Siluro-Devonian System was reported also from the Outer Zone of Southwest Japan and from the Hida mountains, Encrinuridae, Cheiruridae and Scutelluidae were discovered in succession. Dechenella and Thysanopeltella from the Kitakami mountainland were described and re-examined. In the meantime, Ryuji ENDO and Eiji MATSUMOTO (1961) published a comprehensive report on Carboniferous to Permian trilobites. Description of Wenlockian Coronocephalus kobayashii from Kyushu (Takashi HAMADA, 1959) became the first report on complete individuals with cephalon, thorex and pygidium. Furthermore, Motome HIRATA, Jun'ichi HAMADA and T. HAMADA (1967) listed up 9 genera and 12 species of Silurian trilobites from one locality, and this fact must have rectified the past erroneous idea that trilobites were scarce in japan. Since then, data and specimens began to be piled up rapidly.

The fourth period, from 1973 on, is characterized by animated discussions on geologic ages and phylogeny of trilobites referring to the specimens hitherto described. "Silurian Trilobites of japan" by Teiichi KOBAYASHI and T. HAMADA (1974) was the first monograph ever published in japan on japanese trilobites. As many as 40 species were described in the monograph. Thus, japan has come to occupy a high place in the circumPacific region, as well as in Asia, as regards the occurrence of trilobites. Materials of Devonian trilobites also have been successively gained from the Hida

mount:lins (OKAZAKI, 1974; OKAZAKI, TANAKA and TANAKA, 1975; KOBAYASHI and HAMADA, 1975). The results have been compiled into the second monograph by KOBA Y ASH! and HAMADA entitled "Devonian Trilobites of japan" (1975, MS).

On the other hand, study of foreign specimens by Japanese began with the Cambrian trilobites from Northeast China that were reported by KOBA Y ASH! (1930) and R. ENDO (1932). Chinese specimens were studied further by SAITO (1933), ENDO (1935, 1939), and ENDO and RESSER (1939). Before long, KOBA Y ASH! set about his energetic researches on trilobites of the East Asian continent. Since the prewar days KOBA Y ASH! had been pursuing the Cambrian to Ordovician trilobite assemblages not only in China but also in the Korean Peninsula, and published many new views on higher taxa of trilobites. As early as 1935, he distinguished 4 stocks, Agnostida, Redlichida, Corynexochida and Ptychoparida, and discussed fundamental problems in taxonomy of trilobites. His conception is basically taken up in " Treatise of Invertebrate Paleontology, pt. 0, 1959 ". Ontogenesis of Redlichia and Blackwelderia were explicitly studied by KOBA Y ASH! and his collaborator (1951).

These taxonomic studies of trilobites were based on the descriptions of Cambrian-Ordovician specimens not only from the Chinese continent but also from Tasmania, New Zealand, Australia, Yunnan-Tonkin, Kashmir, Siberia, YukonAlaska border, the Canadian Rockies, and the vast circum-Pacific region including Bolivia and Argentina of South America. It is worthy of note that the studies dealt with the materials of the circum-Pacific region with Asia as its center, in spite of the fact that the science of taxonomy had been developed mainly with the


European materials. The above-mentioned far-reaching

studies gave rise, as an inevitable consequence, to discussions of geologic ages, palaeogeography and palaeoenvironment on the basis of trilobite assemblages, and a number of important papers were published with such themes as the Ozarkian problem (1933), the significance of the Masari facies in the Cambrian period (1936, 38), the assumption of the TsinlingKeijo line in the Cambrian-Ordovician period (1930), and the intercontinental sea connections and provincialism (1944, 1965, 1970, 1971, 1972).

One of the remarkable achievements of postwar researches on trilobites outside Japan is the study of geology and palaeontology of Southeast Asia. It began with the work of KOBAYASHI (1956) describing the Cambrian trilobites collected by DEPRAT from the Tonkin-Yunnan border area. In 1957, Upper Cambrian trilobites from the Thai territory of the Malay Peninsula were reported by KoBAYASHI- His report made a great contribution to the geologic history of the older folded mountains of Southeast Asia.

Since then, field survey in Southeast Asia became active and the regional and stratigraphic knowledge of trilobites gained rapidly. In Geology and Palaeontology of Southeast Asia, vol. 15 (1974), as many as 64 species of trilobites were listed up by KOBAYASHI and HAMADA. Among them, 11 genera, 1 subgenus and 36 species are new taxa, including such unique forms as Pseudotrinodus, Langgonia and Prodontochile. The increasing knowledge of these trilobites, along with the progress of the study of Japanese specimens, will throw a fresh light on the older folded region of Asia. The Ordovician-Silurian boundary problem and the early Middle Devonian provincialism were discussed on the basis of trilo-

bites (KOBAYASHI and HAMADA, 1974, 1975).

Crustacea : 1. Conchostraca : The first description of Japanese fossil Conchostraca is found in Matajiro YOKOYAMA (1894). After a blank of 20 odd years, OGAWA and WATANABE (1923) reported 2 species of Mesozoic Estheria from the Korean Peninsula. It was followed by the works of KoBAYASHI and his collaborators (FUJITA, Kmo, T ANI, et a!.) who studied the Mesozoic Conchostraca from Northeast China, Korea and Southwest Japan, over a period of about 10 years. In 1954, KOBAYASHI published a comprehensive paper on Conchostraca of the world, dealing with 48 genera (including subgenera) and 399 species (including taxa of forms below species).

From about 1943, living Conchostraca became the subject of study, and observations and researches were made on the characters of egg and shell, the sexual dimorphism and the relation between climate and life form, in parallel with the study of Mesozoic specimens. The results, compiled by Hisashi KusuMr (1957), contributed a great deal to the study of fossil Conchostraca.

In recent years Mesozoic fossils have been reported from various parts of Southeast Asia. KOBAYASHI (1973, 1975) has pointed out that ancestral forms of Order Conchostraca were sea-dwellers during the Ordovician-Silurian period but afterwards they were all fresh-water forms, and that although they flourished remarkably in the Mesozoic era they vanished entirely in the Tertiary period until they reappear as living forms of today. His indications are important in considering distribution and phylogeny of Conchostraca.

Cambrian Tuzoia (ENDO and RESSER, 1912) from Manchuria and Lower Per-


mian Coreocaris (KOBAYASHI, 1937) from Korea are two rare phyllocariids in East Asia. 2. Ostracoda: The history of Palaeozoic ostracods researches in Japan is still young. The first recorded study is the one by HAMADA (1959) who described and taxonomically examined Devonian "Leperditia" japonica. In the same paper he proposed a new genus Pteroleperditia based on an exhaustive investigation of literature. Kunihiro ISHIZAKI (1963, 1964) detected 7 genera, 18 species of silicified ostracods from Carboniferous limestones. His work was the first faunal description of Japanese Palaeozoic ostracods. He also found a Lower Permian fauna, described 10 genera, 19 species, and established a new genus Hataiella (later emended as Khataiella). From the Middle Permian ISHIZAKI reported 13 genera, 22 species, but the data are too local to be used in discussion of Palaeozoic ostracod faunas in Japan.

From the Sungari Group of Northeast China, 8 genera, 13 species and 1 variety of non-marine ostracods were described by Tetsuro HANAI (1951). This was the first account of Mesozoic ostracods by Japanese, but it was no more than a report of one isolated fauna, as was the case with the Japanese Palaeozoic ostracods.

Cenozoic ostracods of Japan were first reported by Jiro MAKIY AMA (1931) with the specimens from the Pliocene Kakegawa Group, but it was much later that somewhat detailed description was given by ISHIZAKI (1966). Under such circumstances, a series of researches by HANAI, with the results recorded in Studies on the Ostracoda from Japan, I-V, was an epoch-making achievement. Moreover, HANAI (1959) published a bibliography that contains all records of Japanese living and fossil ostracods studies. In

his paper (1970) HANAI discussed schizodont ostracods and readjusted this problematical group, by referring to the carapace and the appendages that are particularly important in living species, as well as to horizons and distributions of fossil species. According to him, schizodont ostracods can be divided into two tribes, Schizocytherini and Paijenborchellini, under Schizocytherinae, in spite of the diversity of their surface ornamentations. 3. Cirripedia: Study of Cirripedia in Japan is represented by a series of excellent works of HIRo (=UCHINOMI) on living species, while records of fossil materials are very few inclusive of some brief accounts on Coronula detached from the skin of whale.

Quite recently, Toshiyuki YAMAGUCHI (1971, 1973) collected and arranged Japanese Tertiary to Quaternary balanids. He re-examined living species and compared them with the corresponding fossil species. In describing 5 species of balanids from the Miocene Mizunami Group, he suggested that fossil balanids can be useful for reconstruction of paleoenvironment because they comprise elements of warm current and those of cold current. 4. Malacostraca: The history of decapods studies in Japan was compiled by Rikizo lMAIZUMI (1963). According to him, the first description of Japanese fossil Crustacea was made by DE HAAN who mentioned Arges parallelus in "Fauna Japonica" (P. F. VON SIEBOLD, 1833-51). Later, Tsune SAKAI (1939) confirmed the occurrence of the same fossil species in Hiejima Island of Kagawa Prefecture.

Mesozoic crabs were recorded very little. There was a report by JIMBO (1894) on Eucorystes japonicus from the Yubari district of Hokkaido. On Ceno-


zoic fossil crabs, IMAIZUMI published his successive studies (1952, 1957, 1959, 1960, 1961, 1962 et seq.). Especially Carcinoplax has been studied in detail.

With regard to fossil shrimp, NAGAO (1931) reported Linuparus japonicus from the Scaphites bed of Ikushumbetsu in Hokkaido. Afterwards, IMAIZUMI (1938) recorded three species of Astacus, late Jurassic fresh-water shrimp, from Lingyuan of Jehol, Northeast China. Records of fossil shrimp are thus scarce including a report by Hiroshi NIINO (1951).

Fossil isopods also are little known. There is only a report by IMAIZUMI (1953) on Bathynomus from Okayama and Ishikawa Prefectures. Anomocardia fossils were recorded in the papers by NAGAO and Kazuo HUZIOKA (1938), NAGAO and 0TATSUME (1938) and NAGAO (1932, 1941) reporting of Callianassa species from the Oligocene to Miocene beds in Hokkaido, by IMAIZUMI (1953, 1957, 1959) on Ctenocheles and Callianassa, and by NISHIKAwA (1972) on unidentified anomocarid crustaceans.

Insecta : The first account of Japanese Mesozoic insects was given by Ienori FUJIY AMA (1973) when he described several species of Triassoblatta, Minesedes and Ominea (Paraplecoptera), Jpsvicioides (Hemiptera) and some species of beetle from the upper Triassic of Yamaguchi Prefecture. Prior to this, Ephemeropsis nymph (mayfly) and Chresmoda (Phasmid) from the late Jurassic of Jehol, Northeast China, had been recorded by Masuzo UENO (1935) and Teizo ESAKI (1945).

Reports on Cenozoic insects have been restricted to those of Neogene Tertiary and younger ages in Japan. IMAMURA (1974) has published a review of the past studies. NAORA (1933) reported some insects of Pleistocene and Miocene.

Lately, FUJIY AMA is devoting himself to description of fossil insects, and through his consecutive studies (1967, 1968, 1969, 1970, 1974) the presence of tropical insects (Heliocopris, etc.) in the middle Miocene has become known. HruRA (1971) also has described a late Miocene carabid.

With regard to Pleistocene insects, there are descriptions of dragonflies by ESAKI and ASAHINA (1957) and ASAHINA (1959), of bugs by OISHI (1931) and HrURA (1961), and of a Papilio and a cicad by FUJIY AMA (1968). Occurrence of fossil insects in various parts of the country has been reported by KINUGASA (1974). Quite recently, nearly 50 families of insects contained in Pleistocene amber at Mizunami have been listed up (H!URA and MIYATAKE, 1974 et seq.).

Other Arthropods: Lower Permian Euproops (KOBAYASHI, 1933) from the Jido Series in Korea is a sole representative of fossil Merostomata in East Asia.

Fossil arachnids had been reported by KISHIDA as early as 1920, but no definite description of specimen was made until a Pliocene spider from Hyogo Prefecture was described by ARIT A and Y AMANA (1970). From amber in the Mizunami district, 11 species belonging in Acarina and 6 species in Arenea have been recorded (H!URA and MIYATAKE, 1974). These specimens, along with those from Shiobara (Tokio SHIKAMA 1965 ; Takashi HAMADA, 1975), await detailed paleontological studies.

On Myriapoda, too, fossil records are scanty. SHIKAMA (1965) reported Diplopoda from the Pleistocene cave deposits in Ehime Prefecture, and H. MINATO (1974) found Rhipidopeltis sp. in amber of Mizunami. No other reports worthwhile mentioning have been published up to now.


Echinodermata-History of Research

Akira MORISHITA*

Echinoids constitute a greater part of fossil Echinodermata studied in Japan, and they are mostly of Cenozoic age. It is strange that no comprehensive reports were written on crinoids in spite of their abundant occurrence in the PermoCarboniferous system throughout the country. There are several papers describing crinoids, namely, by lchiro HAY ASAKA (1924) on '' Actinocrinus" global us from the Carboniferous Omi limestone of Niigata Prefecture, by Hisakatsu Y ABE and Toshio SUGIYAMA (1934) and by Masao MINA TO (1951) on Actinocrinus and Amphoracrinus from the Carboniferous system of the Kitakami mountainland, by Teiichi KOBA YASi:II (1935) on the Jurassic Pentacrinus in Toyoura-gun of Yamaguchi Prefecture and the Jurassic Pseudosaccocoma from Sakawa of Kochi Prefecture. Blastoid was recorded only once when MINATO (1951) reported Nymphaeoblastus from the Carboniferous system of the Kitakami mountainland. Miocene beds in variotfs districts are known to contain some asteroids, such as Luidia and Astropecten, with Amphiura and other ophiuroids, but no scientific description. of these fossils ·seems to have been attempted so far. A· noticeable achievement was the study ·of recent species of ophiuroids . by Hikoshichiro MATSUMOTO (1917) who turned from the zoology course at the Tokyo Imperial

* Institute· of. Earth Sciences, Faculty of Science, Nagoya University, Nagoya.

University to the paleontology course at the Tohoku Imperial University. He described 88 genera, 232 species of ophiuroids in the seas around Japan and indicated their distribution.

Numerous papers were written on fossil echinoids whose abundance ranks probably next to fossil molluscs. The first record was made by NAUMANN and NEUMA YR (1890) in describing Cidaris spines from Sakawa, Kochi Prefecture_ Kotora JIMBO (1894) described Ananchytinarum from the Cretaceous of Hokkaido. This was the first account of fossil echinoids by Japanese. Since then, fossil echinoids were dealt with by many authors, namely, Shigeyasu TOKUNAGA (formerly YOSHIWARA), Takumi NAGAO, Syozo NISIY AMA (formerly AOKI), Yanosuke 0TUKA, lchiro HAY ASAKA, Akira MORISHITA, Masao MINATO, Wataru HASHIMOTO, Matsutaro SHIBATA, Keisaku TANAKA, Masahiro OKUBO, Hiroshi OzAKI, Toshio SAITO, Hiroshi UJIIE and Toshio NISHIO.

The paper by TOKUNAGA (1903) is important in that it drew up, for the first time, a comprehensive list of Japanese fossil echinoids of various geologic ages. He described 25 species in 19· genera of echinoids ranging in age from Cretaceous to Pleistocene. Several years ago, at the British' Museum (Natural History)" in London I found that one specimen of Astriclypeus integer, from the Miocene series of Yamanashi Prefecture, is accompanied by a note, dated

Trans. Proc. Palaeont. Soc. Japan, N. S., (lOOs), .1976


July 1899, sent from TOKUNAGA to Dr. F.A. BATHER, an authority on echinoids. The note reminded me of TOKUNAGA's devotion to fossil echinoids. It is interesting that TOKUNAGA and MATSUMOTO, both distinguished experts on mammals, produced achievements as pioneers in echinoderms also.

A series of studies on fossil and recent echinoids of Taiwan by HAYASAKA (1947-1948, partly under joint authorship with MORISHITA) should be put on record. Since 1950, MORISHITA has been describing fossil echinoids, mostly of Tertiary age, from various parts of japan. In 1960 he fixed their biostratigraphic positions and listed up 30 genera, 65 species and 4 subspecies. He also discussed their relation with lithology (1965) and phylogeny of some of them (1963, 1964).

After all. things taken together, it is undeniable that S. NISIY AMA is the representative of echinoids researchers; His consecutive studies, dating from 1935, dealt mostly with specimens from northern regions, such as the Tohoku district, Hokkaido and Sakhalin, but some from the Kanto district, Southwest japan and

the Bonin Islands were also studied. NISIY AMA's results and the results of all other authors were compiled into " the Echinoid Fauna from Japan and Adjacent Regions, Parts 1, 2 (Palaeontological Society of japan, Special Paper, no. 11, 1966, no. 13, 1968)", which Japan can boast to the world as the most authentic edition on echinoderms. In this edition NISIYAMA gave detailed descriptions of fossil echinoids, amounting to 68 genera, 114 species and 11 subspecies (Paleozoic : 1 genus, 2 species, Mesozoic: 17 genera, 23 species, Cenozoic : 66 genera, 89 species, 11 subspecies), and discussed the characteristics of echinoid fauna of fossil and recent species.

However, paleoecology and evolution of echinoids have been studied very little in Japan. For future studies in these fields, many stratigraphic problems have to be solved first. For instance, Miocene Astriclypeus, Echinolampas, Brissopsis and Kewia should be duly evaluated as significant and effective index fossils. Also, extensive studies of other echinoderms of abundant occurrence, such as Paleozoic crinoids and Tertiary asteroids, are required,

A Short History of Vertebrate Paleontology in Japan

Tadao KAMEl*

The japanese Islands, unlike the continental regions, are scarce of terrestrial formations such as fresh water deposits. Accordingly, fossils of land vertebrates are extremely poor and most of them are allochthonous. They are usually frag-

* Department of Geology and Mineralogy, College of Science, Kyoto University, Kyoto.

mentary and isolated, having been transported over a long distance to be buried in sediments. Moreover, they are often deformed or broken by compression and/ or shearing stress caused by intensive tectonic movements.

The scarcity of fossil vertebrates results in the relative paucity of researchers and research reports, which accounts



for the small rate of vertebrate paleontology among the paleontological researches in Japan. Nevertheless, Japanese vertebrate paleontology had held an internationally important position, based on the geographical uniqueness of the Japanese Islands, being located along the border of the East Asian continent, and has developed in its own way dealing with meager materials. The history of vertebrate paleontology in japan can be divided into five stages (KAMEr, 1968), but the present article reviews the history in four periods by relating it to the worldwide development of paleontology and characteristic progress in japan. The Initial Stage (1868-1900) : This period is characterized by the research work of foreign scientists. It was the period of description. Before the Meiji era, vertebrate fossils were mostly treated as medicines due to the influence of the Chinese thought, or as objects of collection by amateurs or as treasures of temples and shrines, seldom as the objects of modern paleontology.

P.F. VON SIEBOLD, a German doctor in the employment of the Dutch East India Company, came to Japan in 1823 and stayed until 1829. During his stay he introduced japanese fauna and flora to Europe. A fossil elephant that was heaved up from the bottom of the Seto Inland Sea in a fishing net was brought by him to his country. This specimen was described later by K. MARTIN (1887). Prior to this, British paleontologist A. L. ADAMS (1868) reported a fossil elephant discovered in an area " between Edo and Kanagawa ".

During this period, H. FALCONER and R. LYDEKKER studied the Cenozoic mammalian fossils from the Siwalik Hills of India, and the vertebrate fossils from China were introduced for the first time by R. OWEN (1870). Also, there was the

work of K. MARTIN (1883-90) on the Indonesian vertebrate fossils. Under these circumstances, European researchers were directing their attention to vertebrate fossils in japan.

In 1872-74 the "Nihon Sanbutsu Shi" (Products of Japan) written by K. ITO was published from the Ministry of Education. This book recorded some vertebrate fossils. In 1877 when the Imperial University of Tokyo was established and the Tokyo Museum was opened, fossil specimens were gathered and the situation for research activity was improved. Japanese mammalian fossils were described by E. NAUMANN (1882) and D. BRAUNS (1887), both German scientists staying in Japan in those days, although there was a large divergence of views between the two on specific identification, dating and paleoclimate. At any rate, it was in this period that the study of fossil proboscideans became an important theme of vertebrate paleontology in Japan.

The Pioneering Period (1900-1920): Shigeyasu TOKUNAGA and Hikoshichiro MATSUMOTO left their footmarks as Japanese researchers and as pioneers in vertebrate paleontology of Japan. Both of them were zoologists, in contrast to the fact that Matajiro YoKOYAMA and Hisakatsu Y ABE, pioneers in invertebrate paleontology, were geologists.

ToKUNAGA, collaborating with Juzo IWASAKI, described the fossil skull of a large mammal which had been discovered at Togari of Mizunami City, Gifu Prefecture, and assigned it to a proboscidean. This was the first discovery of a skull of Desmostylus (YOSHIW ARA [ = TOKUNAGA] and IWASAKI, 1902). Later, the specimen's similarity to Desmostylus hespents MARSH of California, North America, was noted and it was recorded as


D. japonicus TOKUNAGA and IWASAKI (1914). Before this, desmostylid molars had been found in Shimane Prefecture and Gifu Prefecture in about 1898, but the study of desmostylids that are peculiar to the North Pacific region became an important theme of Japanese vertebrate paleontology for the first time in this period.

On the other hand, the extremely rich contents of the fossil vertebrate fauna of the Chinese continent were being disclosed through the works of R. OWEN, E. KOKEN, M. SCHLOSSER, 0. ZDANSKY and others over a period, from the 1870's to 1903. In 1903, Takuji OGAwA made geological survey of Shantung and recorded several mammalian fossils. H. MATSUMOTO studied mammalian fossils that had been collected from Si-chuan and Honan. He pointed out that the Mammalian fauna of Wan-xian in Sichuan Province comprises two horizons, " upper Pliocene" and " lower Pleistocene". This was the first study which referred to the geologic age of the Wanxian fauna (H. MATSUMOTO, 1915a, b). In the study of japanese fossil vertebrates, it is important to consider them in connection with those of East Asia. This was done by H. MATSUMOTO (1915-1930) who consciously dealt with proboscideans and artiodactyls, particularly bovine and cervine fossils.

Thus, in this period the research field of Japanese vertebrate paleontology expanded remarkably, with various subjects such as fossil proboscideans, desmostylids, artiodactyls-especially cervine fossils-, and the faunal changes of the vertebrates in Japan in relation to East Asia.

The Summarizing and Developing Period (1920-1945): Up to this time most of researches were individual descriptions of

fossils, but during this period a new direction of study came out ; on the basis of hitherto accumulated materials the taxa and their phylogeny were discussed. With such tendency of research activity, faunal descriptions were made and positioning of faunas in the East Asian regions was attempted. Furthermore, in japanese vertebrate paleontology which had been strongly bound with geology was born an attitude turning toward interdisciplinary researches to work in concert with biology, medical science and dentistry.

Till then, the Imperial University of Tokyo where S. To'K:uNAGA taught and the Tohoku Imperial University with H. MATSUMOTO had been the center of vertebrate paleontology in japan, but as time lapsed into this period fossil vertebrates began to be studied also at the Kyoto Imperial University, Hokkaido Imperial University and Waseda University.

H. MATSUMOTO published his views on classification and phylogeny of fossil proboscideans in a series of works with japanese specimens including Moeritherium and Palaeomastodon (1923, 1924a, b, c, 1926a, b, 1929a, b). Jiro MAKIY AMA of the Kyoto Imperial University published a monograph (1938) on the fossil proboscideans of the japanese Islands and neighboring areas, mainly on the basis of the material obtained from the bottom of the Seto Inland Sea. The results of these leading japanese paleontologists were largely adopted. in " Proboscidea, Vols. I, II, 1938-41" by H. F. OSBORN who contributed to the study of proboscideans of the world. It must be mentioned here, however, that in classifying proboscideans MAKIY AMA was a Jumper in contrast to H. MATSUMOTO who was a splitter.

Through the study of desmostylids, a


large quantities of materials were obtained from various parts of japan and from Sakhalin as well. An epoch-making event was the discovery of a complete skeleton at Keton of Sakhalin in 1933. In those days views were diverse on the taxonomic po3ition of desmostylid, but the morphological study in japan was initiated by H. MATSUMOTO (1918a). Late, Takumi NAGAO of the Ho:(kaido Imperial University studied the morphology of Desmostylus teeth and its geological significance (1935a, b, 1977a-d, 1941). Shoji IJIRI worked on the dental morphology and the histology of Desmostylus (1937a, b, 1938a, b, 1939a, b, 1940). And yet, description and restoration of the Keton specimen remained undone.

Study of japanese cervine fossils began with the work of H. MATSUMOTO (1918b, 1926c), which was succeeded by S. TOKUNAGA and Fuyuji T AKAI (1936, 1939), Tokio SHIKAMA (1936a, b, 1937, 1938), and 25 species under 11 genera described so far were summarized by SHIKAMA (1941).

During this period, excavation and investigation of vertebrate fossils were carried out on a large scope in various districts of East Asia and Southeast Asia. Especially the discovery of " Sinanthropus" at Choukoutien in 1926 markedly accelerated the progress of vertebrate paleontology in the Chinese continent. Chinese vertebrate paleontology was conducted chiefly by the Cenozoic Research Laboratory in Peking, which was established in 1927, and by the Geological Survey of China. Besides Chinese researchers, a large number of western scientists took part in the research work.

Japanese researchers also contributed to the development of vertebrate paleontology through their scientific activities in Northeast China (Manchuria), Korean Peninsula and Taiwan. For example, the First Scientific Expedition to Manchou-

kuo conducted excavation at Kuhsiangtung of South Manchuria in 1931. They also carried out excavation at Tamalukou of South Manchuria and at Dokantin of North Korea, disclosing the Late Pleistocene mammalian fauna (TOKUNAGA and Nobuo NAORA, 1939). Ryuji ENDO and T. SHIKAMA, working at the Central Natural Museum of Manchoukuo that was established in 1938, proceeded with their study of fossil vertebrates. Noteworthy achievements were the study of the Djalainor skull (ENDO, 1945) and the discovery of Endotherium (SHIKAMA, 1947), the oldest placental mammal, from the Husin coal formation (Late Jurassic to early Cretaceous). The study of the Eocene mammals from the Hosan Coal Fields of North Korea (SHIKAMA, 1943; T AKAI, 1945) was important in the relation with the Paleogene mammalian fauna of Japan, as observed in the Ube and lshikari Coal Fields. The mammalian fossils of the Ryukyu Islands were studied, too (TOKUNAGA, 1937, 1940; TOKUNAGA and TAKA!, 1939).

With regard to Reptilia, researches were made on such specimens as Monjurosuchus and jeholosauripus (ENDO and SHIKAMA, 1942; SHIKAMA, 1942) and Teilhardosaurus (SHIKAMA, 1947). In South Sakhalin a complete skeleton of Nipponosaurus, a kind of land dinosaur Trachodon, was found and described (NAGAO, 1936, 1938) though its restoration was not performed.

Fossil fishes from the Mesozoic of Manchuria were described by T AKAI (1941, 1942, 1943).

As specimens and information of East Asian vertebrate fossils accumulated, it became necessary to review and summarize the existing materials. It was done especially by comparing them with those of japan and by clarifying the position of japanese materials (T AKAI,


1938, 1939, 1941 ; SHIKAMA, 1941, 1942; NAORA, 1943). It is interesting that at about . the same time the. mammalian fossils from the Chinese continent and aqjacent regions were compiled by Teilhard DE CHARDIN and P. LEROY into a list entitled " Chinese Fossil Mammals" (1942). The list also introduced Japanese specimens fairly extensively.

Mentioned so far are the development and the summary of the four major themes of vertebrate paleontology in Japan, that were presented in the preceding period. But several new themes were indicated in this period.

In the regions where marine deposits are developed well, like the Japanese Islands, study of marine fossil vertebrates is important. Japan abounds in cetacean and pinnipedian fossils, not counting desmostylids. Fossil Cetacea was studied first by MATSUMOTO (1917, 1926a, b), followed by the works of MAKIY AMA (1936) and NAGAO (1941). SHIKAMA and D.P. DOMMING (1970) reported on a Pliocene Sirenia Hydrodamalis which suggests a relation to North America.

Then, differing from the past way of individual description of fossils, a new trend of study came out, as represented by the description of the regional fauna as a whole and its paleoecological study. Along this line there was the work of SHIKAMA on fossiliferous fissure deposits of the Kuzuu Limestone. He made the first report in 1937, but publication of the comprehensive report had to wait until the termination of the Second World War (SHIKAMA, 1949). His work was the first step toward the study of the Japanese Quaternary vertebrates, in particular the abundant fossils in cave deposits, and it was important also in the respect that it presented a problem of the domain bordered by poleontology, anthropology and archeology.

Shoji IJIRI, since 1936, has been conducting histological and physiological researches on the teeth of Desmostylus and proboscidean fossils. Up to that time, vertebrate paleontology in Japan had been concerned mostly with description of specimens from the biostratigraphical viewpoint alone, but IJIRI's work helped to develop methodology which, by deliberating adopting biological techniques, synthesized the biological and geological methods (IJIRI, 1949).

Another advance was the approach to the science of evolution from the field of paleogeography. At this stage the regional biogeography developed into the historical biogeography. !; Biogeography of Japanese Islands " by Mitoshi TOKUDA (1941) exerted a large influence upon geology, paleontology, and the science of evolution, as well as upon biogeography. The correlation of mammalian faunas between East Asia and other continents since the Mesozoic time was discussed by SHIKAMA (1943) and TAKA! (1952).

Period of Expansion and Differentiation (from 1950 on) : Intensification of the Second World War and the chaos of postwar days deteriorated the condition of academic work and caused a temporary paralysis of research activities. The characteristics of the postwar research work are, group studies, large scale excavations and investigations, comprehensive studies in various fields, and activation of international scientific interchanges.

In 1950, the Desmostylus Research Committee (DEREC) was organized, the main object of which was the study of the skeleton of Desmostylus mirabilis NAGAO that had been found at Keton of Sakhalin and preserved at Hokkaido University. With H. Y ABE as the chairman, S. IJIRI, M. MINATO, T. SHIKAMA and F.


TAKA! proceeded with the study, which was intended for development of one major theme of japanese vertebrate paleontology. It happened in the same year that a complete skeleton of Paleoparadoxia tabatai (TOKUNAGA) belonging to Desmostylia was discovered at Toki City of Gifu Prefecture. Part of the research result on this specimen was published (IJIRI and KAMEl, 1961 ; SHIKAMA, 1966).

Accompanying engineering works and constructions that were being carried out on a large scale, findings of fossil proboscideans were frequently reported from various parts of the country. Using these specimens, a systematic study was undertaken on Elephas (Palaeoloxodon) naumanni MAK., a Pleistocene elephant commonly found in japan. Yoshikazu HASEGAWA (1972) studied the innumerable specimens (Takao collection) obtained mostly from the -bottom off Shodo Island in Seto Island Sea, and revealed their individual variation. The specimens from the bottom of Lake Nojiri in Nagano Prefecture and Churui of Hokkaido were studied by Tadao KAMEl and Biroyuki TARUNO (1971, 1973).

While the stratigraphical and chronological studies of the Japanese Quaternary were in progress, the biostratigraphical study based on mammalian fossils, mainly proboscideans, was conducted (KAMEl, 1962; KAMEl and SETOGUCHI, 1970 ; SHIKAMA et. al., 1973).

With regard to fossil cervids, early Pleistocene specimens obtained mainly from the Kuchinotsu formation of Kyushu were studied by Hiroyuki OTSUKA (1965, 1966, 1967, 1968, 1970, 1972). Also, there are the works on late Pleistocene Megaloceros and Alces (SHIKAMA, 1958, 1962; MATSUMOTO et al., 1959; KAMEl, 1958; HASEGAWA, 1968 ; Shingo 0NODERA, 1970).

A significant development during this period was the study of Quaternary mammalian fossils in cave and fissure deposits. It began with SHIKAMA's "The Kuzuu Ossuaries" (1949), but NAORA (1954) also made a comprehensive study on specimens from various parts of the country. Later, the presence of rich fauna was reported from Same Cave in Shiga Prefecture (SHIKAMA et al., 1952), from Mikatagahara in Shizuoka Prefecture (SHIKAMA et al., 1955), from Akiyoshidai in Yamaguchi Prefecture (SHIKAMA and 0KAFUJI, 1957, 1958), from Kumaishi Cave in Gifu Prefecture (OKUMURA, 1973), and from Okinawa-jima and Miyako- jima (HASEGAwA. et al., 1973). The analysis of fauna in these cave deposits offered much information on smaller fossils, such as insectivores and primates (HASEGAWA, 1957 ; SHIKAMA and HASEGAWA, 1958; HASEGAWA et al., 1968; IWAMOTO and HASEGAWA, 1972; IWAMOTO and TAKA!, 1972). This made it possible to study micro-mammals which had been considered a difficult task for the japanese vertebrate paleontologists. · The above-mentioned research works let to synthetic studies connected with anthropology and archeology. Such studies resulted in the discovery of human bones of late Pleistocene age from the limestone area of the Tokai district and from the fissure deposits in limestone of Okinawa-jima. These bones, though fragmentary, were named Mikkabi Man, Hamakita Man, Ushikawa Man and Minatogawa Man, according to their localities (Hisashi SUZUKI, 1962 ; T AKAI, 1966; Naotsune WATANABE, 1973). An early discovery of fossil man was made in 1931 when N. NAORA found a hipbone at the Akashi coast of Hyogo Prefecture. The bone was assigned to early Pleistocene age (Kotondo HASEBE, 1956), but its exact horizon remained unconfirmed.

56 MATSUMOTO, T. et al., edit.: .

An example of such joint researches by anthropologists, vertebrate paleontologists and geologists in Japan is seen in the study of the " Amud Man" and coexisting fauna of the Middle and Near East (SUZUKI and TAKA!, ed., 1970).

In 1949 it was disclosed at Iwajuku of Gumma Prefecture that paleolithic culture had existed in Japan. Since that time, vertebrate paleontologists and archeologists kept on working together, as exemplified by the excavation at Hanaizumj of Iwate Prefecture (1958-60) (MA TSUMOTO et a!., 1959; Kanto Loam Research Group, 1961). Above all, the Lake Nojiri Excavation is unique. From 1962, a large number of people, not only specialists but also non-specialists, have jointly worked in the excavation at Lake Nojiri in Nagano Prefecture. It was the first case in Japan that along with numerous bones of Paleolithic game animals like as elephant and deer, stone implements and artifacts were unearthed from the latest Pleistocene deposits. In many respects, it may be appreciated that this excavation opens the way for new advancement in paleontological, archaeological and geological sciences (Lake Nojiri Exc. Rec. Group, 1975).

The histological and physiological study of fossil teeth, that was initiated by S. IJIRI, was further developed, and with the progress of paleobioche_mistry such modern techniques as X-ray.analysis and electron microscopy were employed. There came out the reports on crystallographic study of fossil teeth (SATO, Suoo and IJIRI, 1957; IJIRI, 1955), on organic matter in fossil teeth (IJIRI and Takayo FUJIWARA, 1958, 1959; IJIRI and KOBAYASHI, 1960), on microstructure of fossil teeth (IJIRI and KAMEl, 1961 ; KAIBARA, 1968; Masahiko AKIYAMA et a!., 1968; KOZAWA, 1974a, b).

On other vertebrate fossils than mam-

mals, reports ar~ still few. Little study was made. on Aves. On Reptilia, however, post-Cretaceous chelonians were found and studied (SHIKAMA, 1953, 1956, 1964; URATA, 1968; OTSUKA, 1970). Ancient ichthyopterygia was recorded from the Lower Triassic beds in Miyagi Prefecture (SHIKAMA, KAMEl and MURATA, 1972), and Jurassic and. Cretaceous specimens are known to occur in other parts of Japan. The Upper Cretaceous formation at Futaba, Fukushima Prefecture, yielded a complete skeleton of plesiosaur (Ikuwo 0BATA, HASEGAWA and SUZUKI, 1970), and a pterosaur was reported from Hokkaido (OBATA, HASEGAWA and OTSUKA, 1972). In Fukui Prefecture, Tedorosaurus of Lacertilia was found from the Late Jurassic beds (SHIKAMA, 1969). Fossil reptiles of younger age are represented by Pleistocene Ophidia and crocodile (SHIKAMA, 1949; SHIKAMA and Goro OKAFUJI, 1958). A gigantic crocodile, 8 m long Tomistoma, came from the Osaka group (Nobuo KoBATAKE and KAMEl, 1966).

Amphibian materials are extremely scanty in Japan ; hitherto recorded are no more than several specimens of Rana, etc., collected from Pleistocene lacustrine deposits and cave deposits (OKADA, 1937; SHIKAMA, 1949, 1955 ; SHIKAMA and 0KAFUJI, 1958; SHIKAMA and HASEGAWA, 1962).

Study of fossil Pisces developed remarkably in recent years. The oldest specimen of Chondrichthyes is the Permian Helicoprion from Gumma Prefecture. With regard to Elasmobranchii, numerous teeth are known to occur in post-Cretaceous beds. Masatoshi GOTO (1972) published a comprehensive report on these specimens. Osteichthyes were recorded in JaRDIN's "Description of New Fossil Fish from Japan" (1919), but at present the Cenozoic fishes are the


major subject of research work. The Miocene marine fauna in the Tohoku and Kanto districts were studied by Jiro SA TO (1962, 1965, 1966). Yoshiro TOMODA et al. (1973) described the Miocene fauna of Iki Island. On the Miocene-Pleistocene freshwater fishes, Teruya UYENO and others made extensive researches, dealing mainly with Cypriniformis of North America and Japan (UYENO, 1961, 1963, 1967 ; UYENO et al., 1962, 1963, 1965, 1975). Also, th~re are the reports on otolith and

fish scale (HAT AI, 1900; 0HE, 1970, 1973; 0HE and ARAKI, 1973; TAN AKA et al., 1966).

Thus, vertebrate paleontological studies in Japan at the present time are manysided. To serve as a window for the internal and international scientific exchanges, the Association for Vertebrate Paleontology of Japan (A VP J) was set up in 1972 within the Geological Society of Japan.

Problematica and Trace Fossils

Hiroshi NODA *

-... -.,., '

This article gives an outline of the history of researches in Japan on problematica and trace fossils which are the records of life of ancient organisms but whose original forms or formative mechanism remain unravelled. The fundamental problems and basic concept on the systematic classification and nomenclature of problematica and trace fossils have been discussed recently by FREY (1973), SIMPSON (1975) and others.** The studies in Japan have been developed under a strong influence of European researchers. At about the time when paleoecology began to be actively discussed in Japan, two remarkable paleoecological books of A. SEILACHER (1953) and R. Th. HECKER (1957) were translated into Japanese by Yoshio KASENO (1957a, b) and by Takeo ICHIKAWA and

* Department of Earth Sciences, Tsukuba University.

** In order to systematize the classification on an international scale, efforts are being made by R. W. FREY and others, including Kotora HATAI of Japan.

Yukio KuwANO (1959). These books contributed to heighten the paleoecological significance of problematica and trace fossils. Particularly the systematic researches on molluscan and crustacean burrowings, excluding sporadic records as "sand-pipe" (JyUzo ARAI, 1955; ARAKI, 1959; HAT AI, 1938-1972 ; KIKUCHI, 1966-67; Koichiro MASUDA and TAKEZAWA, 1961; MASUDA and Hiroshi NODA, 1969 ; SAITO, 1962, etc.), have developed into the field of paleoecological study (ITOIGAWA, 1963a, b, 1974; MASUDA, 1968, 1971a, b, 1972; Tsutomu UTASHIRO, 1968; UTASHIRO and HORII, 1965a, b). Some of the results were good enough for discussing the phylogeny (Kazuo OHSHIMA, 1967, 1968).

The "Trace Fossils and Problematica" by W. HA.NTZSCHEL and " Worms " by B.F. HOWELL, published in the Treatise on Invertebrate Paleontology, Pt. W (1962), gave a strong stimulus to the studies in Japan (for example, Jiro KATTO's work, 1960-1975, on trace fossils from the Cretaceous to Paleogene sys-



terns of the Shimanto Terrain in Shikoku and from the Miocene Series of Wakayama Prefecture, and Keisaku TANAKA's work, 1970, on those from the Cretaceous system of the Ikushunbetsu district. Hokkaido). In the Treatise, HA.NTZSCHEL (1962) re-examined Manchuriophycus (Ryuji ENDO, 1933), Notaculites (Teiichi KOBAYASHI, 1945) and Magarikune (Masao MINATO and Kunio SUYAMA, 1949) that were the forerunning results on Japanese "trace fossils".

As Ichir6 HAYASAKA (1957, 1960) pointed out the ichnological researcher is required to possess knowledge of living forms and to conduct careful observation of the tidal zone for understanding the taxonomical problems of these fossils. In comparison with the European study on these fossils contributing a great deal to reconstruction of paleoenvironment and to paleogeographical analysis, the Japanese one produced few reports and is still backward in the line, although some noticeable results have been pub-

lished by HAT AI and his collaborators, who studied problematical fossils ranging from the Permian (HATAI and MURATA, 1971; HATAI, Masafumi MURATA and KAWAKAMI, 1972; HAT AI, Tamio KOTAKA and NonA, 1972) to the Jomon age (HATAI, 1970), with a view that the fossils would be significant enough if they were distinguishable by morphological characters and were effectively used in geology. They are in continuous pursuit of paleoecological-paleoenvironmental signficance of problematical fossils.

In contrast with the European new method which introduced functional elements into the conventional morphological classification, HATAI and NODA (1975) showed a new approach to the study of trace fossils using Terebellina. It ts very desirable for the propagation of these paleoichnological fossils that they are introduced much more in paleontological texts or illustrated books (KATTO, 1973a, b, 1974; Tokio SHIKAMA, 1975; UTASHIRO, 1971).

Paleozoic Plants

Kazuo ASAMA *

Before World War II, few reports were made on Paleozoic plants of Japan. Japanese paleobotanists were dealing mainly with the Upper Paleozoic plants from Korea and China. It was only after the war that Devonian plants were discovered in I wate Prefecture, followed by the discovery of well-preserved Permian plants at Maiya of Miyagi Prefecture. These findings proved that Paleozoic


plants are distributed in Japan. Later, Devonian plants were reported also from Kochi Prefecture of Shikoku. The presence of Gigantopteris nicotianaefolia and Bicoemplectopteris hallei in the Upper Permian system of Fukushima Prefecture was recorded. Since these species are the representative constituents of the Gigantopteris flora that characterizes the Upper Shihhotse Series in Shansi Province of China, it became evident that the Upper Paleozoic plants in Japan belong to the Gigantopteris flora.



Devonian plants: The first record of Devonian plants in Japan was made by Koichi T ACHIBANA (1950) who described Leptophloeum cfr. australe and Cyclostigma sp. from the Upper Devonian of Iwate Prefecture. Motome HIRATA (1966) reported Leptophloeum rhombicum, Lepidodendropsis sp. and Dometria sp. from Kochi Prefecture, revealing the fact that the distribution of Upper Devonian plants is not limited to the Tohoku district but extends to the Shikoku district in the south. Upper Paleozoic vascular plants: (Lower Carboniferous plants in Malaysia): Lower Carboniferous plants are not known in Japan or Korea. The Lepidodendmpsis flora that was distributed from central China to Yunnan is found in Malaysia, as Lepidodendropsis and Lepidodendron occur at Kuantan of Pahang Province (ASAMA, 1973). (Cathaysia flora in East Asia excluding japan): The name Cathaysia flora was proposed by Thore G. HALLE (1937) for the Carboniferous to Permian plants of East Asia. The Cathaysia flora, in its early stage, was close to the Euramerian flora, but with the passage of time it came to have an East Asian character represented by Gigantopteris nicotianaefolia and Lobatannularia heianensis. Therefore, the later stage of the Cathaysia flora is known by the name GigantoPteris flora. The large simple leaf plant, Gigantopteris nicotianae folia, representative of the Gigantopteris flora, was recorded first from Hunan of China by A. SCHENK (1883). G. nicotianaefolia was described also from Mun-gyong of Korea by Hisakatsu Y ABE (1908). David WHITE (1912) reported G. americana from North America. He considered that this plant had migrated from East Asia into North America via the Bering Strait. Y ABE (1917) discussed the distribution of

Gigantopteris and mentioned that this plant recorded in North America is of Lower Permian age whereas the plant in China and Korea belongs to Lower Triassic. But, it was the voluminous work of HALLE (1927) on the flora of Shansi Province that revealed the whole aspect of the Upper Paleozoic plants in East Asia.

Shigetaro KAwASAKI (1927) established a new genus Lobatannularia which preceded HALLE's Annularites in the date of publication. KAWASAKI gave the full picture of the Heian System of Korea through his studies (1931-1934), partly under joint authorship with Enzo KoN' NO. KAWASAKI's work is comparable to HALLE's study on the flora of central Shansi. KAwASAKI pointed out that the flora of the Kobosan Series, uppermost member of the Heian System, includes Gigantopteris, similar to the flora of the Upper Shihhotse Series of China, and it shows some features common with the Gondwana flora. KoN'No (1960) recorded Schizoneura manchuriensis from the Penchi coal field of northeastern China and showed that the plant, which is undistinguishably similar to Schizoneura of Gondwana in external appearance, was distributed as far north as Manchuria. KoN'NO (1968) also reported the flora that contains Lobatannularia, Gigantopteris and Rhipidopsis at Kaishantun on the eastern border of Northeast China. He considered that the Gondwana elements in China and Korea, such as Schizoneura and Rhipidopsis, migrated from Gondwanaland into Cathaysia land during the Kungro-Kazanian interval (KON'NO, 1960, 1963, 1966).

Genichi KOIDZUMI (1936) maintained that the specimens hitherto reported as Gigantopteris from China, Korea, Sumatra and North America were too much diverse to be included in one genus, and


he divided them into eight genera. Kazuo ASAMA (1959) thought that the

simple leaf Gigantopteris had evolved out of tripinnate frond, and assumed three evolutional series, Emplectopteris series, Emplectopteridium s~ries .. and Kon'noa series. According to.3.his notion, these evolutional series resulted from the change in the environment of North Cathaysia when an oceanic mild climate turned to a continental dry climate. ASAMA (1970) held an opinion that Sphenophyllum in Cathaysia had undergone more advanced evolution than the Euramerian flora and had acquired East Asian characteristics, hence he subdivided the genus into four genera. Also, he ascribed the occurrence of Schizoneura, Sphenophyllum speciosum and Rhipidopsis in Cathaysia and Gondwana to the parallelism of evolution in the two continents, not to their migration from Gondwada to Cathyasia (ASAMA, 1966, 1970).

KON'NO and ASAMA (1963, 1966, 1970)

reported the flora that contains Gigantopteris and Lobatannularia from Thailand· and Malaysia, and indicated that the southern limit of the Gigantopteris flora had extended to Johore. Cathaysia flora in japan : ASAMA (1956, 1967) was the first to record the presence of Permian plants in Japan. He reported Cathaysiopteris, Psygmophyllum, OdontoPteris and Taeniopteris from the Parafusulina Zone at Maiya of Miyagi Prefecture, and revealed that this flora is comparable with the flora of Shansi Series of China. ASAMA and Masafumi MURATA (1974) reported two species of Taeniopteris from Setamai of Iwate Prefecture, and clarified that the Lower Permian flora in Japan consisted chiefly of Taeniopteris.

ASAMA (1974) reported Gigantopteris, Bicoemplectopteris, etc., from the Upper Permian of Takakurayama in Fukushima Prefecture, showing that the representative plants of the Gigantopteris flora existed in Japan, too.

Study of Mesozoic Plants in Japan and Adjacent Regions

Tatsuaki KIMURA*

From 1877 to 1945: In Japan, fossil plants are found in the strata . .of various ages, ranging from Triassic to, ,j~te Cretaceous. According to the present kJIOW:ledge, the Japanese late Jurassic tor:~gply Cretaceous floras are divided in too the Tetori-type flora of the Inner Zone and the Ryoseki-type flora of the Outer Zone.

* Department of Astronomy and Earth Sciences, Tokyo Gakugei University, Tokyo.

The Tetori-type flora resembles the contemporaneous floras in the Siberian palaeofloristic area (V.A. VAKHRAMEEV, 1964, 1966, 1970, 1971), and the Ryosekitype flora is allied to the contemporaneous floras in the Indo-European area, namely, Wealden-type floras (Tatsuaki KIMURA, 1961, 1975; KIMURA and SEKIDO, 1963).

The first researches of the Japanese Mesozoic floras were made by H.T. GEY-



LER (1877) on the Tetori-type flora, and by A.G. NATHORST (1890) on the Ryoseki-type flora. Since that time to the beginning of the 20th century the research work was succeeded by Matajiro YOKOYAMA and Hisakatsu YABE, then resumed by Y ABE's students Seido ENDO, S. ToYAMA, Saburo OISHI and Misaburo SHIMAKURA. Research activities of these palaeontologists covered not only Japan but also China and Korea.

YOKOYAMA described the Tetori-type flora in 1889, the Ryoseki-type flora in 1894, and the late Triassic flora of Yamaguchi Prefecture in 1905. These pioneer works of YoKOYAMA were supplemented by Y ABE. In 1913 Y ABE described the Ryoseki-type flora from Omoto, Iwate Prefecture, and in 1922 he described the Japanese, Chinese and Korean Mesozoic plants preserved at Tohoku University. In 1927 Y ABE published a comprehensive study of the Japanese Cretaceous System, accompanied by description of some Cretaceous plants. Y ABE and TOYAMA (1928) reported early Cretaceous algae from Iwate Prefecture.

Mesozoic plants of Japan are mostly impressions, retaining no texture, except for some specimens from limited localities. M. C. STOPES (1909) and STOPES and K. FUJII (1910) studied the specimens obtained from the Upper Cretaceous System of Hokkaido, and made a histological study for the first time in Japan.

OISHI, working either by himself or in collaboration with Kazuo HuzroKA and Eitaro TAKAHASHI, described the following floras from various parts of the country and largely contributed to the development of the Japanese Mesozoic palaeobotany.

Middle to late Triassic floras: Nariwa flora of Okayama Prefecture (OISHI, 1930, 1931, 1932; OISHI and HUZIOKA, 1935, 1938), floras of Tsubuta, Habu and Mine

Groups of Yamaguchi Prefecture (OISHI, 1932; OISHI and TAKAHASHI, 1936), Shitaka flora of Kyoto Prefecture (OISHI, 1932).

Early Jurassic floras: Kuruma flora of Nagano Prefecture and Toyama Prefecture (OISHI, 1931), flora of the Nishinakayama Formation of Yamaguchi Prefecture (OISHI, 1935).

In addition to the above, they made palaeobotanical studies on Fraxinopsis and Y abeiella (OISHI, 1931), Zamiophyllum (OISHI, 1939), Swedenborgia (OISHI and YAMASHITA [=HUZIOKA], 1935), Die/yozamites (OISHI, 1936), and Dipteridaceae (OISHI and YAMASHITA, 1936). Enzo KoN'No (1944) discussed the phylogeny of Swedenborgia, and HuziOKA (1938, 1939) described Matoniaceous ferns from the Nishinakayama Formation and the lower Monobegawa Group of Kochi Prefecture.

Summarizing the above-mentioned results, OISHI (1940) published his comprehensive paper on the Mesozoic plants of Japan and Korea. He divided the Mesozoic plants of Japan and adjacent regions into Dictyophyllum, Onychiopsis and Angiosperm Series. Chronology of the Japanese Mesozoic plants owes a great deal to Teiichi KOBAYASHI (1938, 1942). In the present paper, the ages of the respective floras are based on the latest results. After the work of ENDO (1925), the

study of the late Cretaceous flora made little progress. Kan KORIBA and Shigeru MIKI (1931) reported Archaeozostera from the Izumi Sandstone. Yuzuru OGURA (1927, 1930, 1931, 1932, 1933, 1941) and SHIMAKURA (1933, 1934, 1935, 1936, 1937, 1939, 1941) largely contributed to the study of Mesozoic fossil wood in Japan. They dealt not only with Japanese specimens but also with materials from China and Korea.


Fossil plants of the adjacent regions became the subject of study in addition to the Japanese ones. YoKOYAMA (1906) studied Jurassic and Cretaceous plants obtained from Yun-nan, Ssu-ch'uang, Chian-hsi, Shan-tnng and Liao-ning of China. Jurassic to early Cretaceous plants were studied also by Y ABE (1908, 1922), Y ABE and ENDO (1934, 1935), Y ABE and OISHI (1928, 1929, 1933, 1938), OISHI (1933, 1935, 1941), TOYAMA and OISHI (1935), OISHI and TAKA HASH I (1938), and KOBAYASHI and YOSHIDA (1944), and the area of study expanded as far as the northeastern region. The histological study of Ginkgoales by OISHI (1933) is related high as a great contribution.

Korea has the Daedong flora ranging in age from Triassic to early Jurassic, the early Cretaceous Nagdong (formerly Naktong) flora, and the late Cretaceous Silla flora. The Daedong flora was studied by YABE (1922), Shigetaro KAwASAKI (1925, 1926, 1939) and KON'NO (1944), the Nagdong flora by Y ABE (1905, 1922), lwao TATEIWA (1929) and OISHI (1939, 1940).

From 1945 on: After 1945, study of fossil plants by Japanese researchers was discontinued for a while. But before long, research work was started by TAKAHASHI, HUZIOKA, KON'NO, Kazuo ASAMA, Toshitsugu OYAMA, Hidekuni MATSUO, and Kazuo OMURA, M. NISHIDA, K. TAKAHASHI, KIMURA and Shinji SEKIDO, and has been continued up to now.

TAKAHASHI, with Goro 0KAFUJI and other collaborators, studied the midele to late Triassic flora of Yamaguchi Prefecture and the Jurassic flora in the Higashinagano, Nishinakayama, Utano and Kiyosue Formations, and clarified the stratigraphic distribution of plant species. He also discussed the post-Mesozoic floral variation in Southwest Japan(TAKAHASHI,

1949, 1950, 1951, 1957, 1967, 1973; TAKAHASH! and G. NAITO, 1950; TAKAHASHI and 0KAFUJI, 1967, 1968, 1970 ; TAKAHASH! et a!., 1965).

KoN'No (1949, 1962, 1968) and KoN'No and NAITO (1960) carried out detailed palaeobotanical studies on fructifications of plants, particularly of Articulates and Coniferales, and opened up many new fields of science. Furthermore, they added new knowledge to the morphology and classification of Dipteridaceae. KoN'No (1972) described Pleuromeia and Neocalamites from the lower Triassic System of Miyagi Prefecture.

The Nariwa flora was added with several genera and species by Nobuo KOBA TAKE (1954), TAKAHASHI (1952) and HUZIOKA (1970).

KIMURA (1959) described the early Jurassic flora discovered in Gumma Prefecture and the early Jurassic flora of Niigata Prefecture, and he discussed distribution, floral composition and characteristics of early Jurassic plants of Japan. In the wake of OISHI (1938), KIMURA discussed the relationship between the Lepidopteris zone and the Thaumatopteris zone (T.M. HARRIS, 1937) in Japan. Kenji KONISHI (1952) discovered and reported the early Jurassic flora in Okayama Prefecture. The Ryoseki-type flora is known to occur in the upper Jurassic System of the Outer Zone of Japan, in the lower Cretaceous Ryoseki Group, and in the lower Monobegawa and upper Monobegawa Groups. The Ryoseki-type plants of late Jurassic age were partly studied by 0Y AMA (1954) and ENDO (1952). ASAMA (1968) discussed the relationship between the evolution of Nilssonia and the climate.

KIMURA, cooperated by Motome HIRATA andY. KANSHA, clarified the stratigraphic distribution of Ryoseki-type plants, and described some new genera and species


(KIMURA and HIRATA, 1975). HUZIOKA (1973) reported W eichselia for the first time from Japan.

NISHIDA (1960, 1962, 1965-1973) described the Ryoseki-type plants and as many as 30 species of ferns and wood of Coniferales from the upper Monobegawa Group in the Choshi Peninsula of Chiba Prefecture. Kiyoshi TAKAHASHI (1972) discerned pollen and spore fossils from the upper Monobegawa Group of Iwate Prefecture.

TheTetori-type plants are distributed in the central part of Honshu of the Inner Zone, ranging in age from late Jurassic to early Cretaceous. The Kiyosue flora of Yamaguchi Prefecture is of the Tetori type. The so-called Tetori-type plants distributed in Gifu, Nagano, Toyama, Ishikawa and Fukui Prefectures are divided stratigraphically into four floras, Kuzuryu, Oguchi, Akaiwa and Tamodani, by their compositional characteristics (KIMURA, 1975).

The Kuzuryu flora of late Jurassic age was described by KIMURA (1958, 1959). The Oguchi flora constituting the main part of the so-called Tetori flora, was described in the papers of GEYLER, YOKOYAMA, YABE, and OISHI. MATSUO and OMURA (1968), KIMURA (1961), and KIMURA and SEKIDO (1965-1975) amended past descriptions of the flora, and additionally described many palaeobotanically interesting genera and species. OGURA, KOBAYASHI and Shiro MAEDA (1951) described Xenoxylon. The Oguchi flora and the Akaiwa flora represent the early and the late Neocomian respectively, and their comprehensive study is under way by KIMURA and SEKIDO. The Tamodani flora of late Early Cretaceous age was described by KIMURA (1975).

Late Cretaceous plants including angiosperms are found at Hakobuchi of Hokkaido, Kuji of Iwate Prefecture, Oarai

of Ibaraki Prefecture, Omichidani along the Ishikawa-Fukui prefectural border, upper reaches of the Asuwa River in Fukui Prefecture, Kamogata of Okayama Prefecture, Suritaki of Hiroshima Prefecture, Kotsuki district of Nagasaki Prefecture, and in the Izumi Sandstone as well as in Wakayama, Hyogo and Ehime Prefectures.

MATSUO (1954, 1960, 1962, 1964, 1966, 1970)described the Asuwa and Omichidani floras and part of the plants in the Izumi Sandstone. He also described the distribution and age of the late Cretaceous plants in Japan.

0Y AMA (1958-1963) and 0Y AMA and MATSUO (1964) described the Oarai flora. OYAMA (1962, 1963) also described part of the Kamogata flora.

There are the works of ENDO (1959) on part of the Suritaki flora, of KORIBA and MIKI (1958) on Archaeozostera from the Izumi Sandstone, of T A TEIW A (1933), K. TAKA HASH I (1958) and KON'NO (1962) on the Kotsuki flora. The age of the Oarai and Kotsuki floras is possibly Tertiary.

Besides the above studies, K. TAKAHASH! (1964, 1965, 1967) described the upper Cretaceous pollen-spore flora of Hokkaido, and discussed the distribution of pollen-spore flora in the Aquillapollenites area. A. MIKI (1971, 1972) described the pollen-spore flora in the Kuji Group of Iwate Prefecture.

The object of study was not restricted to Japan but was extended to Southeast Asia, Korea, and even as far as South America.

T. AKAGI (1954) described the late Triassic flora of the Hongay Coal Field. KoN'No and ASAMA described the late Mesozoic flora of Malaya (KoN'No, 1967, 1968; ASAMA, 1974), the early to late Mesozoic floras of Thailand (KoN'No and ASAMA, 1973; ASAMA, 1974), and the late

64 MATSUMOTO; T. et al., edit.:

Triassic flora of Kalimantan, each time in cooperation with the scientists of the respective nations.

KoN'No (1962) described some species and discussed the stratigraphic distribution of the plant species in the Daedong Group. The study of the Daedong

flora is being continued by KIMURA ·and B. K. KIM. KIMURA and C. VERGARA found abudant late Cretaceous plants in Quiriquina Island of Chile (in MAEDA, Takashi HAMADA, Takeshi CHISAKA, KIMURA and H. TATSUKE, 1972).

Cenozoic Plants

Toshimasa T ANAI*

Studies of leaf and seed floras: (The first period) : Study of Cenozoic plants of Japan was initiated by A.G. NATHORST, C. ETTINGSHA USEN and other foreign researchers. NA THORST (1883) studied the Pliocene plants collected from the Mogi coast of Nagasaki City by the Swedish Vega Expedition. This was the first detailed account of Tertiary flora in Eastern Asia, and it attracted attention of European paleobotanists (SAPORTA, 1883, 1884; ETTINGSHAUSEN, 1883). Later, NATHORST (1888) studied the Cenozoic plants collected from various parts of Japan by the staff of ·the Geological Survey of Japan at that time, and arranged them by localities. For a long period since then, these two works of NATHORST have served as a guide to the studies of Japanese Cenozoic plants.

A.N. KRYSHTOFOVICH, who stayed in Japan from 1917 to 1918, examined fossil plants preserved at the Geological Institute of the Tokyo Imperial University and at other institutions, and wrote several reports on them (1918-1930). R. FLORIN re-examined the ·specimens from

* Department of Geology and Mineralogy, Faculty of Science, Hokkaido University, Sapporo.

Mogi, and published a detailed report on them and on the specimens from Amakusa and Hirado of Kyushu and from the environs of Shimonoseki (1920). In those days his work was an advanced study as it not only described the specimens but also discussed paleoclimate by comparing fossils with living vegetation of Japan.

(The second period): Thus, until about 1920, it was by paleobotanists of foreign countries that the Cenozoic plants of Japan were introduced to the world. After 1920, Japanese researchers began to study Mesozoic and Paleozoic plants of Eastern Asia including Japan, but the studies of Cenozoic plants were still much behindhand.

Since 1928 Seido ENDO published the results of his studies, mostly descriptive works, on Cenozoic plants, and revealed a large number of extinct plants that had existed in the Tertiary period of Japan, Korea and Manchuria. Especially his discussion of paleoclimate (1935) based on the comparison of the Pleistocene flora of Shiobara, Tochigi Prefecfure, with living forest composition indicated a direction of studies of Cenozoic floras. Though ENDO dealt with fossil plants



from various parts of Eastern Asia, many of his papers did not go beyond giving lists of the floral components, excepting the case of the Shiobara flora (1934, 1940). This may account, at least partly, for the retardation of Cenozoic plants studies in spite of the usefulness of fossil plants for Cenozoic stratigraphy. Enzo KoN'No (1931) made a detailed study of late Neogene flora of the central area of Nagano Prefecture, and compared the flora with known Tertiary floras of Japan. It is a matter for regret that his paper gave only illustrations of fossil species without description. H. MORITA (1931-1933) and Haruo 0KUTSU (1933-1943), collaborating with ENDO, continued to describe important species of Cenozoic plants.

In the meantime, Shigeru MIKI started his studies of late Tertiary to Quaternary seed floras (1933 -1941), thus opening up a field that had remained unstudied in Japan till then. His work ushered in a new phase in the paleobotany ·of Japan, and also gave a strong stimulus to the studies of Quaternary geology. Through precise examination of fossil materials MIKI (1941) established a new genus Metasequoia. Later, this unique tree was found growing in the mountains of Szuchuan (Szechwan)" Province of China (H. Hu, 1948), and the fact attracted attE~n

tion of botanists of the world as a " living fossil".

Tertiary plant-bearing beds are typically devloped in northern Japan. Saburo OISHI in collaboration with Kazuo HUZIOKA carried out the biostratigraphic study of the Tertiary system it Hokkaido and Sakhalin, and examined important Tertiary genera one after another (1941-1943, 1954). He also reviewed "Arctic Miocene Flora", the classical work of 0. HEER (1868-1883), and pointed out the confusion in many respects of fossil hori-

zon and identification (1943). During the period from 1930 to 1943,

Cenozoic floras were reported in succession not only from Japan but also from various parts of Korea, Manchuria and Sakhalin. But many of the reports were mere descriptions of important species, and few papers dealt with details of floral composition, excepting the work of MIKI, so that the Cenozoic floral sequence was still unestablished. There were only brief summaries of Cenozoic flora by ENDO (1931) and Y ABE and ENDO (1940).

(The third period): World War II caused a temporary paralysis of research activities in paleobotany. It was after 1950 that new researchers were brought up and the Tertiary stratigraphy of Japan was actively studied. Consequently, researches on· Cenozoic plants came to flourish rapidly.

MIKI (1948) published a comprehensive report on seed floras of Pliocene and younger ages in the Kinki district and surrounding areas, and discussed the floral changes in relation to paleoclimate. Since 1950 he engaged himself in precise examination of these fossils by family, and in clarifying the relations between the distribution of fossil species and living species. His results, though not entirely flawless ·from the biostratigraphic viewpoint, were at a high level paleobotanically and served as valuable references for the studies of late Cenozoic seed floras of Japan. It was also in this period that the first atlas of fossil plant·s in Japan was published by OISHI (1950), to be followed by another by ENDO (1955). HuziOKA (1951-1955) described numerous fossils of broadleafed plants from the Miocene of Korea. Also he disclosed (1950, 1952) that the "Aniai-type" flora and the " Daijima-type" flora, differing from each other in floral composition and


horizon, had been widely distributed in the Inner zone of northern Japan during the early to middle Miocene. OKUTSU, who was studying the Neogene flora of the Sendai area, gave the details of its composition and the biostratigraphy of the area (1955).

For some time after the war, the leadership of research work was still held by those who had been active in prewar days. From about 1950 new researchers began to make their appearance. Toshimasa TANAI published his descriptive works on Miocene plants (1952-1953), and summarized the history of Cenozoic floral changes in Japan on the basis of published literature (1952). Study of the green tuff region was active in those days, which helped TANAT in compiling the atlas of early to middle Miocene plants (1955). About that time the postwar researchers published their first papers, e. g., Hidekuni MATSUO (1952-1954), Kiyoshi TAKAHASHI (1954), Shohei KOKA W A (1954-1955), Keiji SUZUKI (with HUZIOKA, 1954).

MrKr's comprehensive study kept on expanding and his reports (1956-1961) in which a great many fossil plants were re-examined by each family made a good guide to the study of late Cenozoic plants. ENDO (1961-1964) described Paleogene plants of the lshikari coal field, and Kozo NAGAI (1957) summed up the data on the Paleogene flora of Mt. Ishizuchi, Shikoku.

T ANAl (1956-1961) continued to analyze the biostratigraphy and fossil assemblages of Neogene floras in the principal regions of Japan. Furthermore, he distinguished 6 horizons of Neogene floras throughout the country and compiled the history of the floral changes (1961). Meanwhile, K. SUZUKI (1958-1961) was studying the Neogene flora in Fukushima Prefecture. He divided Neogene floras in the southern part of the Tohoku dis-

trict into 7 zones, and presented the history of the floral changes (1961). Sadamasa. MuRAl (1957-1962) published his detailed reports on the Neogene flora of the Shizukuishi basin in Iwate Prefecture. HUZIOKA in his comprehensive paper (1964) gave a full account of the so-called Aniai-type flora of early Miocene in Honshu.

Thus, the researches on Cenozoic plants made a rapid progr~ss with the specimens mostly from Hokkaido and the Tohoku district where fossiliferous beds are developed well,, and the outline of the Neogene floral sequence became almost definite after 1960. On Paleogene floras, however, many problems remain unsolved. Fossil plants beds, ranging in age from late Pliocene to Quaternary. are well developed in western Japan. MIKI and his collaborators studied these beds in detail and presented the outline of the floral sequence. KOKAWA's work (1958-1963) on the relations between the form of Menyanthes seed and the geologic age, and K. SUZUKI's summary (1962, 1965) of the Quaternary floras of Japan are important contributions to the Cenozoic paleobotany.

(The fourth period) : As the general features of Tertiary floras were revealed step by step, there arose a tendency toward re-examination of the floral sequence that would serve as the standard in Eastern ·Asia, and also towards consideration 9f the history of changes of Tertiary forests in the North Pacific basin by comparing them with those on the west coast of North America. Such comparative study of Tertiary forests, with assistance of Dr. R.W. CHANEY of North America, was conducted in various parts of japan from Hokkaido to Kyushu, and efforts for paleoecological analysis of floral composition were continued.


The first report of the results was published in 1963, which gave detailed accounts of Miocene floras of southwestern Hokkaido (T ANAl and N. SUZUKI), of central Akita Prefecture (HUZIOKA), and of Noto-nakajima, Ishikawa Prefecture (MATSUO). The report disclosed that the forest composition was different, both latitudinally and altitudinally, in middle Miocene time (T ANAl, 1967b). As part of the comparative study, MIKI and KoKA w A (1962) made a comprehensive report on late Cenozoic floras of Kyushu.

The study was further extended to various horizons in many parts of Japan, producing detailed reports on the Miocene-Pliocene floras of northeastern Hokkaido (T ANAl and N. SUZUKI, 1965), on the Miocene floras in the northern part of the Nato Peninsula (Shiro ISHIDA, 1970), and on the Paleogene floras of the Kushiro coal-field, Hokkaido (T ANAl, 1970), the Ube co3.l-field, Yamaguchi Prefecture (HuzroKA and Eitaro TAKAHASHI, 1970), the Takashima coal-field, Kyushu (MATSUO, 1967). These reports were compiled into " Tertiary Floras of japan, vol. 2" (1972). Through the above works the Tertiary floral sequence of japan was established as a standard of Eastern Asia, and the results obtained so far were summed up on the basis of the relations between the floras and the paleogeographical and paleoclimatological changes (T ANAl, 1967a, 1972; TAN AI and HUZIOKA, 1967).

Since the latter half of the 1960's, the study of Tertiary floras turned from the descriptive works to the analysis of vegetation and paleoclimate. Miocene floras were the major subject of research owing to their abundant occurrence. There are the works on Miocene plants from the San'in district (Y. HOJO, 1973), from Hokkaido (TANAI, 1971; TANAI and N. SuzuKI, 1972), from Gumma Prefec-

ture (K. SUZUKI et a!., 1970), from Akita Prefecture (HuzrOKA and Kazuhiko UEMURA, 1973), from the environs of Shimono3eki, Yamaguchi Prefecture (HuzrOKA and TAKAHASHI, 1973), from Yamagata Prefecture (Toru ONOE, 1974), and many other studies. HuzroKA (1972) published a comprehensive report on Miocene floras of Korea.

Study of Paleogene floras is still much behindhand because of the difficulties in specific identification. Nevertheless, there are some reports by MATSUO on the Sakito coal-field, Kyushu (1970), the Kishima coal-field, Kyushu (1971), on Tsushima (1971), and by ENDO on the Ikushumbetsu formation of the Yubari coal-field, Hokkaido (1968).

With the late Pliocene to Quaternary floras, detailed researches are being conducted on the floral composition and the paleoclimate, along with the stratigraphic re-examination of fossil plant-bearing beds. Distinguished reports were made by H. NIREI on the environs of Takatsuki City, Osaka Prefecture (1968), on the environs of Kanazawa City (1969), by SUZUKI and NAKAGAWA on the environs of Tsukahara, Fukushima Prefecture (1971),. by Keisuke KURODA on the Atsumi Peninsula, Aichi Prefecture (1966-1967), and by ONOE on the environs of Ebino City, Miyazaki Prefecture (1971). KoKA w A, succeeding MIKI, continued to work on seed flora and published his detailed descriptions of Holocene floras in the Boso Peninsula and the environs of Hamamatsu City (1964, 1967).

Anatomical studies of fossil wood: Morphological study of Cenozoic woods in japan is not as active as that of fossil leaf and seed, but its history is rather long. It began with the report by K. REISS (1907) of Germany, who described Cretaceous to Tertiary woods that were


collected in the course of mineral resources surveys in various parts of Hokkaido. After that, researches on fossil wood were conducted mainly with Mesozoic specimens. Cenozoic wood was dealt with only in a few descriptive papers, namely, on the" Umoregi" (buried wood) from the environs of Sendai (K. Y ASUI, 1917; M. TAKAMATSU, 1929), on the "Matsu-iwa" (pinetree rock) in the Paleogene coal seams of Kyushu (K. OHARA. 1926; Yuzuru OGURA, 1944), and on Quercinium in the Paleogene of Kyushu (OGURA, 1932).

Since 1930 Misabur5 SHIMAKURA engaged himself in the study of fossil wood from Japan and many parts of Eastern Asia, and published his results in succession (1933-1937). He also summarized the distribution of conifers by geologic ages (1939). Shunji WATARI, who succeeded to the research work of Y. OGURA, published the results of his detailed morphological study of Cenozoic wood (1941-1950) from various parts of the country including Iwate Prefecture (Miocene series), Shimane Prefecture (Miocene series), and the environs of Yokohama (Pleistocene series). He also published a comprehensive report on the middle Miocene wood flora, particularly

dicotyledons, from the districts along the Sea of Japan (1962). His work was an important contribution to the studies of fossil wood.

After the war, studies of Cenozoic wood have produced few conclusive results, excepting W A TARI's work (1966) on the Paleogene Taxodioxylon of Kyushu. However, with the abundant occurrence of Tertiary woods in Japan, lively research activities are expected for the future.

Studies of Cenozoic plants in Japan are heading towards ecological researches by means of analysis of vegetation and paleoclimate, based on precise stratigraphy. The intention of the researchers is to elucidate the history of changes of Cenozoic forests in the Japanese Islands, and to reveal their connection with the composition of living forests in Eastern Asia. Efforts are being made also in pursuit of phyletic evolution of fossil species from the morphological standpoint. The pollen analysis, which has made a remarkable progress these years, is offering very valuable information to macrofossil studies. Joint studies by pollen analysts and macrofossil researchers would prove most effective in elucidating the history of forest changes.

Paleopalynology and its History

Shigemoto TOKUNAGA*

Paleopalynology, a branch of palynology, is the study of fossil pollen and spore. The subjects of paleopalynological study cover the whole fossil plant kindgom,

* Geol. Survey Division, Nihon Hiry6 Co., Ltd., Muromachi 2-1, Chuo.ku, Tokyo.

dealing with pollen and spore produced by fossil plants. Paleopalynological methods comprise pollen analysis, description of specimens, stratigraphic correlation and inference of paleovegetation and paleoenvironment from the specimens studied. In recent years a new area of study, to

Trans. Proc. Palaeont. Soc. Japan, N. S., (100s), 1976


work on physical and chemical properties of pollen and spore, has been opened.

The history of paleopalynology in Japan, in comparison with the development of this field of science in the world, is summarized in four stages, introduction, accumulation of material, analysis, and new technology, as represented by the number of related literature (Table 1).

The introductory stage dates from 1935 when the technique of pollen analysis was introduced into Japan, to be followed by practice of pollen analysis, particularly on pollen from peat beds. Since paleobotany in Japan had been initiated by foreign researchers (G. NATHORST, R. FLORIN, A. N. KRISHTOFOVICH, etc.) and the subject of study was mainly fossil leave3, the start of analytical researches using microfossils (such as diatoms, pollen and spore) was made much later.

The succeeding stage, from 1950 on, is marked by concentration of pollen analysis for the specimens obtained mostly from high moor peat in Honshu. As a result, pollen zones in and after the final glacial stage were established on the basis of the changes in pollen assemblages; then, the last glacial substage was denoted by L and the postglacial stage was divided into RI, RII and Rill (Jun NAKAMURA and Matsuo TSUKADA, 1952-1974). There was another opinion in which the whole Quaternary system is divided into different pollen zones, A to F (Norio Fun, 1970). The both divisions were based on the transition of paleovegetation that reflects changes in the paleoenvironment. Therefore, it may be defined that the Japanese paleopalynology enteted the period of material accumulation.

In the wake of pollen analysis of the Quaternary deposits, analysis of the Tertiary deposits, especially lignite and coal, was commenced. Its initial object was

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correlation of coal seams. The fine correlation of Paleozoic coal seams by means of fossil spore and the success of palynological investigations of lignite or brown coal fields in western countries gave a strong stimulus to Japanese palynologists, and the palynological characteristics of coal seams in the principal coal fields of Japan were disclosed (Kiyoshi T AKAHASHI, Y oshio OKAZAKI, Seiji SA TO, Shigemoto TOKUNAGA, and others). And yet, the material available for setting up representative pollen indices for the respective coal seams was insufficient. In the meantime, there developed two trends of research work, one was intended for correlation, by means of morphological discrimination and description of fossils, from a purely practical standpoint, and the other conformed to the nomenclature of botanical classification. In either way, regional pollen diagrams of Japan were compiled, but the arrangement of all available data is not complete yet for general correlation on a more extensive scale.

It was in 1964 that the presence of fossil spore in Mesozoic deposits of Japan was reported for the first time. Pollen assemblages in the Omine area where the Triassic system is distributed and in the Tetori area of the Jurassic terrain are too poor to deserve description. Up to now, occurrence of well-preserved Cretaceous specimens is limited to the Hokkaido and northern areas of the Tohoku district. Lately, good assemblages have been found in the lower Cretaceous system of Iwate Prefecture (T. TAKAHASHI, 1974). The oldest fossils are the ones reported from the Permian system of Kyoto Prefecture, but they are meager for stratigraphic use (K. TAKAHASHI, 1969).

The scarcity of Mesozoic specimens in Japan may be ascribed to tectonic movements that could have affected the strata and destroyed fossils, or perhaps the plants

that produce pollen and spore were few. Through the analysis of pollen and

spore assemblages the changes in regional flora of early Tertiary and younger ages are being clarified. In pollen analysis the knowledge of the East Asian floral region is especially important. In Asia, differing from Europe where Tertiary pollen assemblages that indicate significant changes, a weighty clue would be found in the changes of pollen of plants that grew in temperate climate.

Stratigraphic horizon of the Liquidambar-Nyssa-Carya pollen assemblage in the Tertiary system of Japan tends to move upward as the latitude of localities becomes lower, which brings about a new interpretation of correlation by means of fossil plants.

In the Mesozoic pollen and spore, however, many common genera are found including Aquilapollenites which is characteristic to northern regions such as Alaska and Siberia. Therefore, the Mesozoic specimens must be studied from a global standpoint.

For higher precision of analysis, paleontologists' interest in microstructure of fossils is deepening, and the use of phase microscope, electron microscope and scanning electron microscope is becoming popular.

In recent years, the degree and nature of alteration of pollen, spore and other parts of plant have come to attract attention of researchers because these are useful as the elements for measurement of geothermal temperature. Thus, a new area of study is being developed.

From the standpoint of oil prospecting in Japan, dating by the color of fossil pollen and spore is under investigation. No conclusive result has been produced as yet, on account of the fact that the age of subject beds is limited to Tertiary and Mesozoic. At any rate, very precise


analysis is required to attain the object. Future development of paleopalynology

in Japan wili be many-sided, for it comprises various areas of study, namely, microstructure of specimens, their physical and chemical properties, as well as elucidation of paleoclimate and paleoen-

vironment. In comparison with western countries, the research work in japan is concerned mainly with relatively younger formations, and so the paleopalynology in japan and Southeast Asia is expected to make a unique development.

Algal Conservatism

-Symbiosis between earth science and biology-

Kenji KoNISHI*

The history of the researches on fossil calcareous algae in japan can be divided into three periods.

The first period : The first report on algal fossils was made by Kyugaku NISHIW ADA (1897), who studied the reefy Megami-Ogami Limestone of Shizuoka Prefecture in his graduation work at the Geological Institute, Imperial University of Tokyo, and identified the species to "Lithothamnium ramossissimum REuss" known from Leithakalk of the Vienna basin. In Europe at that. time, palaeobotanist RoTHPLETZ (1891) discovered the living fossil Archaeolithothamnium, and divided Corallinaceae into the major groups, Lithophyllum- and Lithothamnium-, by the difference in conceptacles of asexual reproductive organ. His paper was an impact to botanist contemporaries FoSLIE and HEYDRICH.

Several years later, algal nodules from the Riukiu Limestone of Shuri and from limestone lenses in the Yaeyama group of Iriomote-jima, Okinawa Prefecture, were sent to the specialists in Europe

* Department of Earth Sciences, Faculty of Sciences, Kanazawa University, Kanazawa.

through Bunjiro KoTo, and were identified as Lithothamniscum ( =Lithophyllum) nahaense (sp. nov.)' and "Lithothamnium ramossissimum" by HEYDRICH (1900) and NEWTON and HOLLAND (1902), respectively. This was the period when many important papers were written on the taxonomy of living Corallinaceae, inclusive of those by Kichisaburo YENDO (1902-1905).

Hisakatsu Y ABE, who long since realized the importance of calcareous algae as limestone-forming organisms, studied in Europe before he took his chair with the newly founded Tohoku Imperial University. While entrusting KARPINSKY (1910) of St. Petersburg with the study of Mizzia velebitana and Stol/eyella ( =Mizzia) yabei collected at Mino Akasaka of Gifu Prefecture, Y ABE himself (1912) was dealing with Solenoporaceae, discussing their phylogeny and classification. In this study, he described Metasolenopora ( =Solenopora) rothpletzi (sp. nov.) from the Torinosu limestone at Sakawa, Kochi Prefecture, and Petrophyton miyakoense (gen. et sp. nov.) from the Miyako formation of Iwate Prefecture which were the first new taxa of fossil



algae established by Japanese. The second period : Y ABE's constant

interest in calcareous algae was manifested in his comprehensive report (1922) on localities and horizons of Tertiary coralline algae in various parts of the country. Before long, specialists in fossil algae appeared among his students. The first of them was Riuji ENDO (1924) who made a biostratigraphic study of the Carboniferous-Permian System of Hikoroichi, Iwate Prefecture, as his graduation work, and discovered Permian Chlorophyta. Besides ubiquitous Permian iV!izzia (HAY ASAKA, 1922; 0ZA W A, 1924), the species recorded by ENDO included three genera of Alpine Triassic Dasycladaceae, differing from the results of PIA (1912, 1920) who was undoubtedly the world authority on the subject in those days. For this reason, and also with the cautious attitude of Y ABE (e.g. 1952), ENDO confined himself to give just an outline of his result.

In the wake of ENDO, Y ABE asked Shiro TOYAMA to elaborate the algal study of the Jurassic Torinosu Limestone. As the results, several new genera and species of Late Mesozoic calcareous algae were described (Y ABE and ToYAMA, 1928), and it was confirmed later that these algae were cosmopolitan reef-associated species. In 1932 TOYAMA met with an accidental death, but his follow-up report was published in 1949 as a joint work with Y ABE, and the study of the Torinosu calcareous algae was succeeded by his fellow student Rikizo IMAIZUMI (1965).

Adopting the systematics of Mme LEMOINE (1911-17, and later) based on vegetative rather than reproductive tissue, Wataru ISHIJIMA (1933-1954, and later) began to specialize in the taxonomy of Corallinaceae and vigorously described and classified the important species of Cenozoic coralline algae of japan and

Taiwan, along with re-examination of the specimens recorded by Y ABE. He (1943) also proposed a new Cretaceous genus of possibly Chlorophycean affinity from the Miyako Formation.

Characteristic of this period was that a large number of japanese geologists engaged themselves in the geological survey of the Chinese continent. Occurrence of stromatolites of algal origin from the early Palaeozoic beds and the Sinian (Chentan) System was reported by Y ABE, Kin-emon OZAKI, Teiichi KOBAYASHI, Susumu MATSUSHITA, ENDO, Rinji SAITO, Hiroshi OZAKI and Eitaro TAKAHASHI. Possibility of their serving as index fossils of the Precambrian biostratigraphy was suggested by KOBAYASHI (1933). It was unfortunate that stromatolites were erroneously assigned to an inorganic origin when the algal nature of "Manchuriophycus" ENDO (1933) was rejected by the nominator himself (1966). Stromatolites of the Chinese continent have large potentiality to become a treasury of Precambrian microfossils of Asia. A few examples of fossil stromatolites have been reported from Japan (KONISHI, 1959 ; KONISHI and OMURA, 1965).

The third period : With the termination of the war, ENDO returned to Japan and in 1949 he resumed the biostratigraphic study of Palaeozoic calcareous algae. His energetic activity continued until 1969 when he died of illness. While accomplishing his toilsome duties as the Dean of Faculty of Science and Literture, Saitama University, and later as the University's President, he published 30-odd papers (1951-1969) on calcareous algae, proposed more than 10 new genera, and discussed phylogeny of Dasycladaceae and other taxa. His descriptions included species from the Philippines and Thailand. Our knowledge of Carboniferous to


jurassic calcareous algae of Far East owes a greg_t deal to his results. His work has been succeeded by his students Mankichi HORIGUCHI (1957, 1962, 1965) and Manjiro NAKAMURA (1971).

ISHIJIMA also was rep:ltriated from Taiwan, and compiled the results hitherto obtained into a monograph which was published in 1954. His three texts (1933, 1950, 1956) on Japanese fossil algae are among the few guidebooks written in japanese, inclusive of the atlas of fossil plants by ENDO (1966). ISHIJIMA's steady research work is revealing the characters of fossil florules (mostly Corallinaceae) from the Philippines, Pakistan and the Galapagos Islands, as well as from japan and Taiwan.

In 1953 the Japanese Society of Phycolqgy was established under the lead of Yukio YAMADA assisted by TOKIDA, TAN AKA, SEGA W A, ARASAKI, IMAHORI, HIROSE, and others, and this gave rise to the insistence to base the studies of fossil algae on the knowledge of living algae in Japan (KONISHI, 1954, 1961). At the 79th meeting of the Palaeontological Society of Japan in the autumn of 1961, a symposium on fossil algae was held attended by phycologists of living algae.

One of the traits of this period was that the studies of fossil calcareous algae became active in North America, largely owing to the efforts of ]. H. jOHNSON, REZAK, and WRAY. jOHNSON visited algal herberia of japanese universities in connection with his monographic work on the Pacific corallines from 1947 to 1953, and published papers on fossil algae from Ishigaki and Kitadaito, Okinawa Prefecture (1961, 1964). Also, there was a strong tide among the American palaeontogists to be inclined toward ecology and biogeography (NEWELL, LADD, CLOUD). The studies of calcareous algae accelerated this trend

(e. g., jOHNSON and KO:'-IISHI, 1956-59; KONISHI and EPIS, 1960). At the same time, the role of calcareous algae as rock-forming organisms in depositing carbonates was revalued (e.g., LOWENSTAM, GINSBURG), after the prominent pioneering work by PIA (1926), and calcareous alg3.e came to occupy an important part of carbonate sedimentology. Mineralogical and geochemical examinations of hard tissue of calc1reous algae and other calcifying organisms, along their morphologies, are now essential means for facies analysis of carbonate deposits and for restoration of palaeoenvironment and its diagenetic history. In Japan, this type of research is being conducted lately (KONISHI et al., 1968 and later) and will be concentrated toward the palaeoclimatic as well as eustatic studies of the sea floor sediments. Finally, study of Coccolithophorids, the calcareous phytoplankton, has begun in japan and is producing biostratigraphic results from both subaerial and submarine sections (Toshiaki TAKAYAMA, 1967 and Shiro NISHIDA, 1969).

japan is blessed with abundant deposits of algal origin, such as the shallow-water carbonates, ranging from subtropical reef complex to subarctic "maerl" pavement, and the "high temperature type" stromatolite that forms hot spring deposits. These valuable deposits await future sedimentological study, in which a palaeontological approach is indispensable.

Palaeontology as well as her offspring, palaeophycology will continue to develop, by occasionally shifting the position of its center of gravity but basically maintaining once established symbiosis, that is mutualism, between earth science and biology. This prospect is strongly suggested by the retrospect of the japanese researches on fossil calcareous algae.


Concluding Remarks

Tatsuro MATSUMOTO

In concluding the concise history of palaeontology in japan, I should like to give short remarks of my own view instead of a well balanced summary of the preceding chapters.

Scientific works on fossils from Japan started about a century ago, when reconnaissance geological survey was undertaken in several areas of Japan. A hundred years history of palaeontology in japan may be divided into the following four stages, depending much on the activity by generations and also on world development of palaeontology : (1) Stage I : Period of pioneers' works (1873-1911) (2) Stage II : Period of works by a smaller number of specialists (1912-50), with two substages divided by the establishment of the Palaeontological Society of japan (1935) (3) Stage III: Period of increasing number of research works in various fields, with more frequent international exchange of knowledge (1951-1970 on) (4) Stage IV: Toward renewed palaeobiological works (1971 on) These stages may overlap to some extent in actual research works and the dates indicated in parentheses are conventional, although the boundary date marks the commencement of something new (see chronological table).

Stage I. Our hundred years history is much shorter than the history of palaeontology in certain countries in Europe. They have a cultural background of


Renaissance, piles of natural history collections and voluminous monographs by generations.

Despite these handicaps, our predecessors, especially the late Professors Matajiro YOKOYAMA and Hisakatsu Y ABE, father and mother of the palaeontology in Japan, among others, endeavoured to learn the up-to-date palaeontology from Europe, especially from eminent palaeontologists in Germany and Austria in that period, and themselves accomplished valuable research works on various kinds of fossils obtained by geological reconnaissances.

In this stage, which corresponded to the period of descriptive works of classical monographs in the history of world palaeontology, some other monographic descriptions were given on fossils from Japan by European authors, e.g. GEYLER, NAUMANN, NEUMA YR, MOJSISOVICS, NATHORST, FLORIN etc., as well as by several other Japanese, e. g. Kotora ]IMBO+, Gordon YAMAKAWA+, Kono YAsm+, Shigeyasu YOSHIW ARA + [ =TOKUNAGA] etc.

Stage II. The beginning of the second stage may be marked by the establisment of the palaeontological institute in Tohoku University (Sendai) (1912) under the leadership of Prof. Y ABE, where palaeontologists were educated and then specialized respectively in the study of a particular taxonomic group or a particular geological era, e. g. Ichiro HAY ASAKA in Palaeozoic corals and brachiopoda, ShOshiro HANZA w A in larger

+ deceased



foraminifera, Takumi NAGAo+ in Terthry and Cretaceous mollusca, Saburo SHIMIzu+ in Mesozoic ammonites, Seido ENno+ in Cenozoic plants, Ryilji ENno+ in trilobites and c1lcareous algae, Shichihei NOMURA in Cenozoic mollusca, Haruyoshi FUJIMOT.) [ =HUZIMOTO] in fusulina, Saburo OISHI+ in Mesozoic floras, Toshio SUGIYAMA+ in rugo::;a, tabulata and strom:1toporoids, Kin'emon OZAKI in early Palaeozoic fo3sils, Motoki EGUCHI in Mesozoic and Cenozoic corals, Syozo NISIY AMA + in echinoids, Wataru ISH !JIM A in calcareous algae, Misaburo SHIMAKURA in fossil wood:;, Kiyoshi ASANO in smaller foraminifera, Kotora HATAI in brachiopoda, Tokio SHIKAMA in vertebrates and so on. Meanwhile eminent students graduated from the University of Tokyo were like-wise specialized in various fields, e. g. S. TOKUNAGA+ in mammals, Shingo YEHARA + in Me3ozoic faunas, Jiro MAKIY AMA in Cenozoic mollusca and other faunas, Enzo KoN'No in Palaeozoic and Early Mesozoic palaeobotany, Yoshiaki OZAWA+ in Late Palaeozoic fusulina and other faunas, Teiichi KoBA Y ASH! in Lower Palaeozoic cephalopod3, trilobites, etc., Yanosuke OTUKA + in Cenozoic mollusca, Fuyuji TAKA! in vertebrates and so on.

Geological departments weree3tablished in 1921 in the University of Kyoto, in 1930 in Hokkaido Univ. (Sapporo), in 1939 in Kyushu Univ. (Fukuoka), in 1941 in Tokyo Univ. Education, in 1943 in Hiroshima Univ. and so on. More palaeontologists of younger generations, thus, started to work, but their numbers were by no means enough. Although the Palaeontological Society of Japan was established in 1935, it was still a branch of the Geological Society of japan for the financial reason (i. e. too small number of members).

To sum up, the second stage can be

defined as a period of works by a comparatively smaller number or specialists. Their activities were, however, remarkable and the materials for the study were not only confined to japan proper but also expanded to the geologically related neighbouring areas.

Stage I I I. The third stage started when Japan was recovering from the severe damage by World War II. In 1951 the Palaeontological Society of japan issued No. 1 of the New Series of their journal and also No. 1 of their Special Papers. As President of the Society Prof. Y ABE and then Prof. T. KOBA Y ASH! took leadership in this rising period.

Owing to a new policy of educational system, universities and other institutions (including museums) have been renovated and increased in .number. People have got more freedom of doing what they wish and palaeontology has become more popular. Thus in the third stage not only many of the specialists who were active in the second stage continued to work but also numerous students of younger generations participated in palaeontological studies. The fields of activity have become more diverse and extensive, including pollen dispersed by winds and nannofossils from submarine areas.

Being aided by air transportation and affected by the issue of such comprehensive works as the Treatise, Osnovy, etc., and also increased number of journals, the international exchange of knowledge has become much easier and more frequent than in the preceding stages. Economic improvement and peaceful state in japan have been favourable for sound development of sciences. Under these circumstances the palaeontological achievement for these 25 years can be said remarkable, publishing various kinds of papers in great numbers. Thus, the Palaeontological Society of japan issued


No. 100 of the new series of their quarterly jounal at the end of 1975 and will issue No. 20 of their Special Papers in 1976. There are several other special or regional or temporary reaearch groups which have given rise to or are going to give rise to interesting results.

It should be noted that before the issue of the Society's journal the restlts of palaeontological researches in Japan had been mainly published by the universities, thoug:1 at irreguhr. intervals, aided by governmental financial support. This system was followed even after the establishment of the Palaeontological Society. Thus, the volume number of the "journal", "Science Reports", or "Memoirs" of several universities in jap::m have exceeded 20 or even 40. The National Science Museum and several other museums have their own publications in which palaeontology is included, whereas the Geological Survey of japan has no series of palaeontological papers, except for some special volumes. "Geology and Palaeontology of Southeast Asia" (edited by T. KOBAYASHI and Ryii.z6 TORIYAMA or Wataru HASHIMOTO) has been published since 1964 up to vol. 16 in 1975, again with grant in aid of the Ministry of Education. There are some palaeontological papers in other journals, e. g. Proc. japan Academy, japan. jour. Geol. & Geogr., jour. Geol. Soc. japan, Mem. Geol. Soc. japan, Kaseki [=Fossils], Chikyu-kagaku [=Earth Sci.], etc. The Pal. Soc. japan published in 1961 a compiled Catalogue of the Type Specimens in Japan and in 1963, A Survey of Fossils from japan illustrated in Classical Monographs (reproducrion of the illustration in classical monographs published before 1900, with revised explanatory notes). In addition to several textbooks (written in japanese) of palaeontology in advanced courses there are three series of Atlas

of japanese Fossils (1970-1976) (Tsukijishokan) with fine illustration and explanatory text.

Despite these active publications important papers have sometimes been missed to be cited by authors of foreign countries. A good bibliography has been and will be published by the Pal. Soc. japan (in SPecial Papers at the interval of 10 or 15 years). Of course some japanese authors have contributed their papers to certain international journals or books.

There have been, thus, considerably voluminous palaeontological works in Japan, but most of them have been concerned with the systematic palaeontology and biostratigraphy, with descriptions of particular taxonomic groups or faunas or floras of particular areas or ages. As the palaeontology was delayed to start in japan, it was natural and unavoidable to stress the systematics and the stratigraphical palaeontology for a considerable period. In fact Japan and her neighbouring areas, situated at palaeobiogeographically important position, are provided with diversity of faunas and floras in various geological ages. Accordingly some of the results have given outstanding contributions to the world problems in these fields of palaeontology, as explained in the foregoing chapters.

This may be the primary reason why the chapters of this concise history are divided in accordance with major taxonomic groups. However, this way of writing would not be favourable for modern palaeontology, in which research projects tend to be focussed on certain palaeobiological subjects of general interests rather than on particular taxonomic groups.

Stage IV. In Japan renewed palaeobiological works have recently been increasing. In this sense the fourth stage

A Concise History of Palreontology in Japan 77

can be said to have started already, overlapping the third.

Yes, even in the systematic palaeontology more biological foundation should have been taken into consideration. In fact in the years as old as the second and third stages there were such works. Hikoshichiro MATSUMOTO's (1923-24) monographs of fossil mammals, and E. KoNNo's palaeobotanical papers (1929-73), Tamio KoT AKA's (1959) monograph on the Cenozoic Turritellidae and Tetsuro HANAI's (1970) paper on some ostracods exhibit successful examples along this line. Similarly MAKIY AMA's (1924) paper on the evolution of Umbonium, KoBAYASHI's (1935, 36, 37, 47) and SHIMIZU's (1934) works on the internal structures of cephalopod shells and also Shoji IJIRI's (1937, 39) attempt in the embryology and the hystology of Desmostylus tooth were outstanding at those dates for the stress to more palaeobiological aspects.

To conclude this concise history let us give short comments on recent activity in japan with prospects for modern palaeontology.

The population concept has proved fundamentally important in modern biology. For some reasons, however, this was delayed to be introduced to the palaeontology in japan. In fact it may be considerably difficult to analyse satisfactorily the populations on fossil materials, but this should never be taken as a reason to reject the population concept from palaeontology. Biometric and statistic analyses of samples of considerable size which were carefully collected from stratigraphically well ordered sequences have recently been undertaken by several authors of younger generations for taxonomic and other purposes, giving rise to interesting results. They are mostly on molluscan fossils, but Tomowo OzAwA (1975) has presented a successful

work on Permian fusulina in tracing the evolutionary change of Lepidolina as well as in proposing a revised classification of the group.

Itaru HAY AMI (1973) studied the intrapopulational variation of a Pliocene-Recent species of scallop, Cryptopecten vesiculosus, from japan, with a very interesting result that samples of later ages include two clearly discrete phenotypes and that the remarkable change in the frequency of phenotypes during geological times can be interpreted as resulting from the accumulation of a mutant gene by natural selection. This way of approach, if adequately combined with genetical experiments, would bring forth renewed contributions to the problem of evolution. Keeping pace with this kind of approach an attempt should be done to bring the biostratigraphic zonation so up-to-date as to meet with the requirements of modern genetics. This has been recently discussed by HAY AMI (1971) and HAY AMI and 0ZA W A (1975). On the basis of numerous specimens

from the successive horizons of the Upper Cretaceous sequences, Kazushige TANABE (1974, 75) attempted to make clear the evolutionary change in functional morphology on bivalve shells of the group of Inoceramus (Sphenoceramus) naumanni and ammonite shells of the species of Scaphites and Otoscaphites, with promising results. This may exemplify another fresh aspect of palaeontology in japan.

In addition to the population concept recent improvements in techniques and methods should be briefly mentioned here. A textbook written in Japanese entitled "Methods for the study of fossils" [=Kaseki no Kenkyu Ho] was published in 1971 (Kyoritsu Pub. Co., Tokyo) by a number of collaboratory authors. Since Susumu HONJO's (1964, 67) pioneer works


at Hokkaido University, quite a number of palaeontological laboratories in Japan have been equipped with scanning electron-microscopes and several universities have computer centres. Cultivation of some animals is carried on at several laboratories for palaeontological purposes as equipments of chemical analyses are used for the same purposes. On the occasion of the annual meeting of our Society in January 1975, a symposium was held on the scanning electron-microscopic study of fossils, including both the technical aspects and the observed results.

Being aided by technical improvement the micro-and ultramicro-structures of "hard parts" (shells, skeletons, bones, teeth, vegetable organs, tissues, etc.) and their ontogeny are being studied in more detail, as exemplified by Iwao KOBAYASHI (1964, 1971), KAIBARA (1968), and Yokichi TAKA Y ANAGI et a!. (1968). This would be concerned with comparative anatomy in more precision and also be useful for deeper understanding of the function of a particular organ or morphological character. Another development from the study of the structural details is in the problem of biomineralization, which in turn is related with the problems of fossilization and diagenesis on one hand and those of biochemistry and accordingly even those of pearl fishery, gnathology and medicine on the other hand.

Organic matter is often pre( er Jed within the microstructure. Biochemical inspection of the organic matter (mainly amino-acids) in some well preserved fossils had been attempted already in 1958 by IJIRI in Japan and further developed by him and his followers. However, this seems to have been rather involved in the calcification and later alteration problem, without giving so far fruitful results in palaeophysiology. Evolutionary

change in proteins, especially that of nucleic acids, may indeed be a fascinating problem, if ever be made clear, but there is much to be done in treating adequately fossil materials for this purpose. Kenji KoNISHI (1971) and a research

group around him endeavour to know the basic factors which control the isotopic abundances in the organic tissues, especially in the shelly part. I hope this approach of isotope-palaeontology would become one of the useful means in palaeobiological studies.

Palaeoenvironmental analyses on fossil biota or individual fossils have developed gradually in Japan. Palaeoclimate, for instance, has been discussed since YoKoYAMA's date based on the faunal or floral analyses of various ages, but the interpretation has recently become to be done on more up-to-date grounds and in mo;e international or global view point. The biogenic calcium carbonates of certain taxonomic groups are being examined for their potentialities as quantitative thermometer or salinometer, whereas the up-to-date sedimentological analyses can provide a useful information on the palaeoenvironment, as remarked by KONISHI and Hisayoshi IGO (1973). Palaeobathymetry is much needed but may be more difficult, if quantitatively required. Attempts have been done by several authors on some Cenozoic mollusca and foraminifera. Katsura OYAMA (1952, 1973 etc.) indicates with abbreviated letters or symbols the vertical distribution and bottom characters of molluscan species on the ground of his expert knowledge of natural history. This is really useful but may be somewhat special. Recent and fossil reef complexes have been actively studied from palaeoecological and other viewpoints as in other countries.

A Concise Histo1·y of Palaeontology in Japan 79

In the proper field of palaeoecology intensive studies have been carried on to analyse fossil assemblages in certain Neogene foraminiferal and molluscan faunas on the basis of association of species and also lithofacies of fossiliferous deposits, as typically exemplified by Hiroshi Unm (1962 et seq.), Tadamichi 0BA (1969), Junji ITOIGAWA (1961), Kiyotaka CHINZEI and Yasuhide IWASAKI (1967). Most of the fossil molluscan assemblages can be regarded as representing the benthic community, judging from the recurrence of the same occurrence. An interesting example of "paralled community" has been shown by their study. The investigators intend to make clear the chronological change of fossil communities which would, according to them, contribute to the problem of evolution. This may be right, but in my opinion, the ecological conditions of a species or better to say those of Mendelian populations should be more directly concerned with adaptation and natural selection in leading trends of evolution. The evolutionary change of the ecosystem would be another important problem in palaeontology and also in historical geology. However, this could never be answered by treating only the material of local biota of a limited time interval.

Tsugio SHUTO (1957) attempted to treat the problem of speciation on palaeoecological factors, but his example could have been a result of biofacies differentiation which was made clear by his own precise faunal analysis (SHUTO, 1961). The results of his study on larval ecology of prosobranch gastropods (SHUTO, 1974) are significant for their biogeographical distribution including the problem of isolation and consequently modes of speciation.

As to the recent advances in biostrati-

graphy and palaebiogeography notes have already given in the foregoing chapters and only additional remarks are given here on a few points. In these fields the international cooperation has been especially emphasized. For instance, the Silurian-Devonian boundary has long been internationally discussed and recommendations were submitted in 1972 by the Committee, with which Takashi HAMADA is incorporated as a member (see HAMADA, 1973). The international Permian-Triassic conference was held at Calgary, 1971, in which biostratigraphic evidence and the problem of organic crisis were contained. Keiji N AKAZA w A, Kametoshi KANMERA and TORIY AMA contributed papers to this conference bringing well compiled data from Japan or eastern Tethys (see LOGAN and HILLS [ed.], 1973). The same problem has been continued to be studied by NAKAZAWA and his coworkers on the materials of southern Asia where international cooperation has been taken. Their results will be published soon.

Another good compilation of biostratigraphic correlation of all the known fossiliferous formations has recently been achieved for Southeast Asia (KOBAYASHI and TORIY AMA [ed.], 1975). Although the correlation is tentative for some parts, the published results mark a milestone for further advances.

An atlas of palaeobiogeography (HALLAM [ed.], 1973) was published with fine international compilation of contributors in which MATSUMOTO took part in the palaeobiogeography of late Cretaceous Ammonoidea.

As one of the projects of IGCP an internationally cooperative work is being performed on the mid-Cretaceous zonation and correlation, with which MATSUMOTO, TAKA Y ANAGI, and others (1975) are incorporated.


An international conference on Pacific Neogene stratigraphy will be held at Tokyo, 1976, where the biostratigraphy will occupy the main part. Combination of biostratigraphic correlation with radiometric dating and also with palaeomagnetic allocation will be one of the interesting points of discussion, as suggested already by Nobuo IKEBE et a!. (1973).

International cooperation has long been taken and is increasingly important in submarine investigations, in which various kinds of microfossils are important for correlation and palaeoenvironmental interpretation. In some cases, however, megafossils were occasionally found in the boring cores.

Recently a renewed significance has been afforded to the regional palaeobiogeography in connexion with the theory of sea-floor spreading and plate tectonics. Although interests of these aspects are rather in historical geology, the palaeobiogeography itself should be essentially related with the problem of speciation and evolution.

The aim of biostratigraphy is also in historical geology, but we need fundamental palaeontological studies on the organic evolution and related subjects for further refinement of the biostratigraphic

zonation and correlation. The above remarks comprise my

personal opinions, and this chapter could be written in more or less different ways by other authors. There may be some other important aspects of the recent palaeontology, particularly micropaleontology, in Japan which I am afraid to have omitted or missed to stress. Anyhow, I believe that sound development of palaeontology in the future several years would depend on the well balanced advances in both fields of palaeobiology and stratigraphical palaeontology, which should be intimately connected with the up-to-date biology and also renewed aspects of geology. In every field the international cooperation and exchange of knowledge would become more and more important in the future. Fresh discoveries and creations, as well as further improvements on the foundation of previous works, would promote the real advances.

Acknowledgements . . Although I myself is responsible for the above remarks, I owe much to a number of persons, especially Dr. T. KOBAYASHI who wrote in Japanese a short historical review ten years ago and has also contributed an introduction to this issue and Dr. K. CHINZEI who helped me in giving upto-date information on palaeoecology.

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CONTENTS


Preface

KoBAYASHI, Teiichi: Introduction to the History of Palaeontology in Japan...... 1

UJIIE, Hiroshi: Paleontology and Society of Japan .. .. .. .. .. .. .. . . .. .. .. .. .. .. .. 10

IGO, Hisayoshi : Paleozoic Microfossils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • 16

TAKA y ANAGI, Y okichi : A Brief History of Post-Paleozoic Micropaleontology . . . . 19

KATO, Makoto : Coelenterates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

SAKAGAMI, Sumio : History of Bryozoological Research . . . . . . . . . . . . . . . . . • . . . . . . . . 29

YANAGIDA, Juichi: Brachiopodology in Japan-A Historical Review • . . . . . . . . . . . . 30

0BAT A, lkuwo : Cephalopods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . • . . . . . . . . . . 33

SHUTO, Tsugio : Fossil Molluscs ...................•....................... ·. . . . . 38

HAMADA, Takashi : Arthropoda • . • . . .. . . . . . . . . • . . . . • • . . . • . . . • . . . • • • . . . • . . . . . . . . . • 44

MORISHITA, Akira: Echinodermata-History of Research ........................ 49

KAMEl, Tadao: A Short History of Vertebrate Paleontology in Japan ........... 50

NODA, Hiroshi: Problematica and Trace Fossils ................................. 57

ASAMA, Kazuo : Paleozoic Plants . • . . . . . . . • . • . . . . . . . . . . . . . . . . . • . • . . . . . . . . • . . . • • . . 58

KIMURA, Tatsuaki: Study of Mesozoic Plants in Japan and Adjacent Regions . . . 60

T ANAl, Toshimasa : Cenozoic Plants ..................... , . . . . . . . . . . . . . . . . . . . . . . 64

TOKUNAGA~: Shigefri~ti): Paleop~lynoloJ; and its History ............. , . . . . . . . . . . 68 ~ ' ..

KoNISHI, Kenji: ·Algal Conservatism-Symbiosis between Earth Science and

·" ·Biology·.: ... ~- ... .-.•.................•.............••.•. ' .......•...........• 71

MATSUM()TO, Tatsuro: Concluding Remarks .................................... 74

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