植物研究雑誌 1. 1. Jpn. Bot. 71: 71: 168-177 (1 996)

A Revisioo 00 the Chioese Megafossils of F agus ()

Yu-Sheng Lm a， Arata MOMOHARA b and Sheng-Wu MEI a

aDepartment aDepartment of Palaeobotany ，Nanjing Institute of Geology and Palaeontology ， Academia Academia Sinica ，Nanjing ，210008 CHINA; bDepartment bDepartment ofEnvironmental Studies ，Chiba University ， 648 Matsudo ，Chiba ， 271 JAPAN

(Received (Received on June 29 ， 1995)

revision A revision on the Chinese Cenozoic megafossils of Fagus is undertaken. Seven species are recognized ， among among which one is represented by cupule fossils ，while all the others are leaves. The oldest species ，F agus sp.1 with with similarities to two Chinese living endemic ， F. engleriana Seem. and F. longipetiolata Seem. ，is documented documented from the Late Eocene to Early Oligocene of Guizhou Province ，S. W. China. This species also seems to to represent the earliest megafossil record of Fagus in the world. In the meantime ， two species previously documented documented as Fagus (e.g. ， F. chinensis Li and F.? feroniae Unger) are now excluded from the genus. The present present study shows that “ F. englerian α"-like leaves were common during the Cenozoic era in China. Furthermore ，the Chinese fossil records appear to display a low diversity of beeches in the geological past as compared compared with the recent Chinese beeches.

F agus is a quite small genus within the family evolutionary trend of F agus around the Pacific Ocean Fagaceae ， but it is frequently encountered in fossil during the Cenozoic. In the present paper ， an attempt floras floras of the N orthern Hemisphere. Modern beeches ， is made to revise the megafossils reported from however ， are now disjunctly restricted to temperate China and then discuss their geological history. zones zones and mountainous regions of the subtropics in the the Northern Hemisphere (Chang and Huang 1988). Materials and Methods China with more than half of the species in the genus Fossil specimens checked by the present authors

F agus ranks as one of the main centers of distribution. 訂 e mainly housed at the N anjing Institute of Geology These Chinese species are now distributed in the and Palaeontology ，Academia Sinica. Those which mountains mountains of southern China (Cao 1995). are not available to us were examined on the basis of More and more paleobotanical works have pro- published accounts. As most of the Chinese materials vided vided details on the fossil Fagus ，particularly from are leaves ， which all are only preserved as impres- Japan ， North America and Europe (e.g. ， Tanai 1974 ， sions ， a foliar architectural analysis on most of extant Zetter Zetter 1984 ， Minaki 1985 ，Kvacekand Walther 1991). Fagus was undertaken for comparisons with fossils. Unfortunately ，no review ofthe Chinese fossil records Cleared leaves were prepared by using a routine has has been carried ou t. Tanai (1974) excluded the Chi- method (Hickey 1973 ， Li 1987). Hickey's terminol- nese nese beech fossils in his comprehensive paper on the ogy on foliar morphology (Hickey 1979) is followed

-168 ー June June 1996 Joumal of Japanese Botany Vo l. 71 No. 3 169

through through the present paper. species of Fagus in China (see Table 1， Figure 1). For Hill Hill and Read (1991) noted that leaf architecture of easier recognition ，a simple key to the species is given modem Nothofagus (Fagaceae) alone has limited first. value value for infrageneric . Liu (in pr~ss a) mentioned a similar case for betulaceous leaves. Our Key to the fossils of the Chinese F agus recent recent study on leaf architecture of F agus shows that 1. Material preserved as leaves it it is also true for F agus although the foliar architec- 2. Laminas big ，and number of secondary veins

tural tural characters between the only two J beeches apanese high (ca. 16) …...・ H ・..…..，・ H ・.....・ H ・.....・ H ・..F. sp. 1 (Fagus crenata B l. and F. japonica Maxim.) are quite 2. Laminas small ，and numbe .r of secondary veins

useful useful for classification. Without strong evidence ，it about 10... ・H ・...... ・ H ・....・ H ・....・ H ・-…...・ H ・-… .F. sp. 2 might be a safe way for current revision that uncertain 2. Laminas in middle size ，number of secondary specific specific names are applied for most cases. veins around 8-13 3. 3. Margin entire to undulate …・

Fossil Fossil Records of F agus in China …...・ H ・...F. engleriana Seem.

About 779 megafossil species belonging to 281 3. Margin se 打 ate genera genera and 99 families ofhigherplants and bryophytes 4. Leaf index low (ca. 187) ，and leaf base cor-

reported reported from the Chinese Cenozoic up to and includ- date ...・ H ・..…...・ H ・.....・ H ・..…… F. praelucida Li ing ing 1993 have been found throughout the country 4. Leaf index high (more than 200) (Li u，Guo and Ferguson ， in press) ， but there are only 5. Margin between adjacent teeth more or less

a few fossils of F αgus documented in the Chinese straight ...・ H ・.F. stuxbergi (Nathorst) Tanai literature. literature. Meanwhile ， these fossils ， mainly leaves ， 5. Margin between adjacent teeth convex .…・

訂 e mostly poorly preserved. Although features of …F. galbanifolia Guo

vegetative vegetative and reproductive organs in F agus are rela- 1. Material preserved as cup 'u les ....・ H ・....・ H ・.F. sp. 3 tively tively stable (Jones 1986) ，misidentifications are still being being made in the Chinese paleobotanicalliterature. 1) Fagus sp. 1，Zhang ， Papers Stra t. Paleon t. 138 ， pl. The following is a brief revision of the seven fossil 1 ，fig. 15 (1 983) [Fig. lA]

Table Table 1. Comparison of megafossil species of F agus in China a

Taxa Taxa Size Number of Living Geological secondary secondary veins equivalent(s) age

F. F. sp. 1 10 cm x 5 cm 16 F. engleri α na/ Late Eocene to F. F. long 伊etiolata Early Oligocene F. F. galh αnifolia 4-6 cm x 1.8-2.2 cm 8-10 F.lucida Oligocene F. F. sp. 2 3 cm X 1. 2 cm 10 F. multinervis/ Oligocene F. F. engleriana F. F. stuxbergi 4-10 cm X 2-5.6 cm 1cト 13 F. engleri αna Early 恥1iocene F. F. engleriana b ヲ 8-10 F. engleriana Miocene F. F. praelucida 5.6 X 3 cm 8-9 F.lucida Pliocene F. F. sp. 3 0.9 cm (l ength) F. engleriana/ Early Pleistocene F.lucid ，α

aAll aAll the taxa are fossilleaves except that F. sp. 3 represents cupules. bThe bThe size of the species is unknown. 170 170 植物研究雑誌第71 巻第3 号 平成 8 年 6 月

Fig. Fig. 1. Selected fossilleaves of F agus from China. A: F agus sp. 1. B: F. sp. 2. C， D: F. galbanifolia Guo. Bar = 1 cm.

Discussion: Discussion: One fossil leaf reported by Zhang is extremely difficult to elucidate which one is cor- (1 983) is actually two broken of portions beechlike rec t. laminas laminas which are overlapped each othe r. Consider- Zhang (1 983) assigned the fossils to the Eocene in ing ing features of margin and secondary veins of the age on basis the ofleaf assemblages and stratigraphical lower lower lamina ， the pronounced zigzag midrib and correlation. Recently ， she thought it might be as late relatively relatively straight secondary veins of the upper leaf ， as the Late Eocene to Early Oligocene (Zhang per- the the two broken leaves should be attributed to the sonal communication). genus genus F agus even though no strong evidence presents. Material: Deposited at the Guizhou Regional Geo- A concept of Tanai' s leaf index (l ength/width x logical Survey. 100) 100) (see Tanai 1974) is employed for identifying Locality and age: Panxian County ，Guizhou Prov- different different species within the genus of F αgus. The index ince ， southwestern China; Late Eocene-Early is is quite simple and convenient to group fossil beech Oligocene. leaves. leaves. For the cu 打 ent fossils ，their estimated leaf 2) Fagus galbanifolia Guo ，Paleon t. Atlas Jilin 571 ， indexes indexes are probably around 200 ， which show the pl. 262 ，figs. 4，8 ，9 (1 992). [Fig. C1 ，D] fossiIs fossiIs might be close to F. engleriana Seem. in the Discussion: The fossilleaves with their relatively “Fagus grand ゆlia group" (see Tanai 1974). How- well preserved venation pattern and distinct tooth ever ，Zhang (1 983) suggested that the fossil leaves morphology indicate that even in the absence of a resembled resembled that of F. longipetiolata Seem. Anyhow ，it cuticle they should be attributed to the genus Fagus. June June 1996 Joumal of Japanese Botany Vo l. 71 No. 3 171

The most evident character is the comparatively big ， it is also possible that both of the fossil species in this asymmetrical asymmetrical teeth with convex/acuminate basal side locality could belong to one and the same species. and straight/concave apical side. This distinct feature However ， in order to substantiate this idea ，more has has not been documented in any fossils from China. material is needed. The present treatment can only be

Therefore ， the specific status of the cu 汀 ent fossil considered as provisiona l. should should be reasonable. It appears that the fossil has entire or undulate Judging Judging from the tooth morphology ， the following margin although Guo (1 992) reported as being ser- extant extant species are probably closed to the present rate. The gross mo 叩hology of the fossil is rather fossils ， those are F. crenat αB， l. (in Japan; only se 町 ate similar to those of young leaves of F. multinervis type) ， F. hayatae Palib. ex Hayata var. zhejiangensis N akai. Furthermore ， the present fossil is also closed to Liu Liu & Wu (in Zhejiang Province ， eastem China) ， F. F. engleriana Seem. in its marginal nature. lucida lucida Rehd. & Wils. (in southem China) ，and F. Material: Housed at the Nanjing Institute of Geol- pashanica pashanica Yang (in Sichuan Province ， southwestem ogy and Palaeontology ，Academia Sinica. China and Zhejiang Province ， eastem China). The Locality and age: Shanhe Town ，Ya 吋i County ， cu 町 ent fossils ，however ， are still far from the above Jilin Province ， northeastem China ， Oligocene (upper mentioned mentioned extant species in overall nature although part of Hunchun Formation). they they are much similar to F. lucida Rehd. & Wils. ， one 4) Fagus stuxbergi (Nathorst) Tanai ，Tanai ，J. Fac. member in the “Fagus longipetiolata group" (see Sc i. Hokkaido Univ. 296 ， p l. 1 ，figs. 4，5 ，8 ， 10; fig.- Tanai Tanai 1974). text 2，g寸(1 976); Li and Yang ，Acta Palaeon t. Sin. The leaf index is ca. 220-270 that shows the fossils 209 ， p l. 2 ，fig. 5; pl. 3 ，fig. 8 (1 984); Guo ，Paleon t. could could be close to those of Tanai' s“Fagus grand ザolia Atlas Jilin 571 ，pl. 263 ，fig. 8; pl. 264 ，figs. 7，8 (1992). group" group" (see Tanai 1974). Tanai (1 974) concluded that F agus palaeocren α ta Okutsu ，Tanai ， Birbal Sahni this this group contains four species ， F. grandifolia Ehrh. ， Ins t. Palaeobo t. Spec. Pub l. 70 ，pl. 4 ，fig. 4; pl. 5 ，figs. F. F. japonica Maxim. ， F. multinervis Nakai and F. 1，2，4，6，7 (1 974). engleriana engleriana Seem. ，which are all with entire to undu- Discussion: In leaf shape ，venation and character- late late leaf margin. From this point of view ，Tanai' s Ieaf istics of the leaf margin ， the cu 汀 ent fossil leaves are index index is problematic as we doubted before. doubtless attributed to F agus although a great varia- Material: Material: Housed at the Nanjing Institute ofGeol- tion in gross mo 叩hology of these leaves occurs. The ogy and palaeontology ，Academia Sinica. total number of specimens in the fossil flora where F. Locality Locality and age: Shanhe Town ，Ya 吋i County ， stuxbergi (Nathorst) Tanai dominates is about 13 ，i.e. Jilin Jilin Province ， northeastem China; Oligocene (upper ca. 16.25% of the whole assemblage. It is further part part of the Hunchun Formation). believed to resemble F agus stuxbergi (N athorst) Tanai 3) 3) Fagus sp. 2，Guo ，Paleon t. Atlas Ji lin 571 ，pl. 269 ， from the Pliocene Mogi flora of Japan (Li and Yang fig. fig. 8 (1992). [Fig. lB] 1984). Discussion: Discussion: The fossilleaf was recovered from the The leaf index of the present fossilleaves is as high same stratum as F agus galbanifolia Guo. They are ， as 210 that agrees with that of the “Fagus gr αndifolia however ，rather different in their tooth morphology group" (see Tanai 1974) and is particularly close to and leaf size. The present fossil has a smaller lamina the species F. engleriana Seem. However ， Li and (see (see Table 1). It seems to be necessary to separate Yang (1984) considered the fossilleaves approximate them as completely distinguishable species. Of course ， those of F. longipetiolata Seem. F agus stuxbergi 172 植物研究雑誌第71 巻第3 号 平成 8 年 6 月

(Nathorst) (Nathorst) Tanai is recently considered as a closely Japanese endemic beech ，Fag lf: s crenata Blume (see similar similar species to the living F. crenata B l. (“ Fagus Kitamura and Murata 1979 ， p. 282; Ohwi 1984 ， p. sylvatica sylvatica group" of Tanai's) in Japan ， F. hayatae 377). Actually ， the fossil is more similar to one ofthe Palib. Palib. ex Hayata (“ F agus sylvatica group" ofTanai' s) Chinese endemic species F. engleriana Seem. than to in in Taiwan and F. pashanica Yang (F agus longipetio- those of F agus crenata Blume in the nature of its leaf lat αgroup" of Tanai's) in central China (Uemura shape ， trend of secondary veins and especially its

1988). 1988). Without anatomical preservation the present undulate to entire margin. However ， the cu 打 entfossil fossilleaves ，it is difficult to interpret whose opinions leaves show some slight differences with those of F. are are correc t. engleri αna Seem. in the number of pairs of secondary Fagus stuxbergi (Nathorst) Tanai from China oc- veins and the size of the lamina. For example ， the curs curs only from one locality (Liu in press b). Li and fossils have fewer secondary veins (about 8-10 pairs) Yang (1 984) considered the flora an early Miocene and smaller leaves. Unfortunately ， the poor preserva- one one based on comparison with Japanese Neogene tion and incomplete fossil material prevented further floras. floras. That is ， the conifers and gymnopserms in the comparisons (e.g. ， anatomical features) from being Chinese Chinese fossil flora display similarities to those in the made. Early Early Miocene Ainoura and Aniai floras (for defini- Material: Depository unclear. tion tion of these floras ， see Tanai 1961). However ， the Locality and age: Weichang County ， Hebei Prov- angiosperms angiosperms are largely different and none of the ince ， northem China; Miocene. species species ofFagaceae ，which are so characteristic of the 6) Fagus praelucida Li， Li and Guo ，Paleon t. Atlas Chinese Chinese flora ，exist in either of these J apanese floras East China 297 ， p l. 139 ，fig. 1 (1 982). (Liu (Liu in press b). In this respect ， the Chinese flora is Discussion: Although only one specimen has been much more similar to the Middle Miocene D 吋ima found ，its leaf shape ，venation and m 訂 gin show it to flora. flora. Thus this so-called early Miocene flora might be a species in F α gus. The fossil is characterized by its eventually eventually require to be transferred to the middle sinuous margin with a few small teeth and 8-9 pairs Miocene. Miocene. More material is needed in order to clarify of secondary veins. Li and Guo (1982) suggested that its its stratigraphical position (Liu in press b). the present fossil resembles F. palaeocrenata Okutsu Material: Material: Housed at the Nanjing Institute ofGeol- (synonym of F. stuxbergi ， see Tanai 1976 ， p. 298 ， ogy and Palaeontology ，Academia Sinica. Uemura 1988 ，p. 119) and F. protojaponica Suzuki Locality Locality and age: Dunhua County ，Jilin Province ， from Japan although their leaf forms are differen t. northeastem northeastem China; early Miocene (Tumenzi Forma- Furthermore ， Li and Guo (1982) considered the Chi- tion). tion). nese fossilleaf more closed to one Chinese endemic 5) 5) Fagus engleriana Seem. beech ， F. lucia αRehd. & Wils. Fagus sieboldii End. ，Depape ，Pub l. Mus. Hoangho The leaf index is about 187 that might suggest that 9 ，fig. 2 a-f. (1 932). the present fossil is related to the third group of Discussion: Discussion: As early as 1932 ，Depape reported a Tanai's “Fagus longipetiolata group". Li and Guo fossil fossil species by using an extant beech species ，F agus (1 982) came to the same conclusion since F. lucida sieboldii sieboldii End. ， based on several Miocene leaf impres- Rehd. & Wils. is concluded in the group by Tanai sions sions from Hebei Province ，northem China ， but no (1 974). As the fossil is the only one recovered and detailed detailed description was provided. F agus sieboldii without any cuticles preserved ，it would be safer to End. End. has long been treated as a synonym of one recognize the present fossil as a provisional species June June 1996 Joumal of Japanese Botany Vo l. 71 No. 3 173 before before more materials are available. the fossi l. It should be mentioned ，however ，that even The fossil is even considered as a representative fossil cupules of Fagus from the Japanese Middle more closely to Fagus hayatae Palb. ex Hayata than Pleistocene (Osaka Group ラ westem Japan) show diι to to F. lucida Rehd. & Wilson by Shen and Boufford ferentmo 叩hology with any living ones (Miki 1933). (1 988). This interpretation might be without any It is possible ，therefore ，that the Chinese cupule fossil evidence. evidence. is not really co 打 esponding to that of F. engleriana

Material: Material: Deposited at the Nanji 時 Institute of Seem. in despite ofthis living beech distributes around Geology and Palaeontology ，Academia Sinica. the fossillocality. Locality Locality and age: Nanfeng County ，Jiangxi Prov- Material: Housed at the Guizhou Regional Geo- ince ， southeastem China; Pliocene. logical Survey. 7) 7) Fagus sp. 3. Locality and age: Panxian County ，Guizhou Prov- F agus engleriana Seem. ，Zhang ，Paleon t. Atlas South- ince ， southwestem China; early Pleistocene. west west China 490 ， p l. 165 ，figs. 9-10 (1 978). Discussion: Discussion: Some early Pleistocene cupules were Chinese Megafossil Excluded from F agus first first reported from Guizhou Province ， southwestem Two more species have been referred to F agus by China China (Zhang 1978). They are unique reproductive various authors. After comparing them with extant fossil fossil organs of F agus recorded in China. They were species of F agus ， the present authors consider them to considered considered to resemble those of F. engleriana Seem. be misidentifications. since since they had a cupule 4-valved with 0.9 cm long 1) Alnus prenepalensis Hu & Chaney ，Hu and Chaney ， lobes lobes (Zhang 1978). The lobes are oval to rounded in Palaeon t. Sin. 30 ，pl. 10 ，figs. 1，4，6 (1940); WGCPC ， shape shape while those ofthe living one are linea r. Further- Cenozoic Plants from China 59 ， p l. 57 ，figs. 1，2，4 more ， the cupules of F. engleri ω1G Seem. have very (1978). F agus chinensis Li ，WGCPC ， Cenozoic Plants distinct distinct appendages on their surface ， which are not from China 46 ， p l. 26 ，fig. 7; pl. 42 ，fig. 5; pl. 44 ，fig. present present on the cu 汀 ent fossil specimens. From this 2，text-fig. 25 (1 978); Tao andDu ，ActaBo t. Sin. 275 ， point point of view ， the fossils resemble those of F. lucida table 1 (1 982). Rehd. Rehd. & Wils. The size of cupule of F. lucia αRehd. Discussion: Three localities from which fossil & Wils. ，however ，is about twice than those of the leaves referred to as Fα gus chinensis Li have been fossils. fossils. According to the photographs provided by recovered are the Fushun Coalmine ， northeastem Zhang (1978) ， the fossil cupules seem to have some China ，Shanwang Diatomite mine ， eastem China and sparsely sparsely distributed round scars that are probably Tengchong County ， southwestern China. This resulted resulted from fossilization. In addition ，no stalks are misidentification was first put forward by Liu (in preserved. preserved. Those taphonomic processes and poor press a). Fossil leaves described as F. chinensis Li preservation preservation make the fossils impossible for further from the first two localities differ considerably from classification. classification. typical leaves of Fαgus ，especially in the secondary As the fossil cupule was not present in collec- the vein trends (see Jones 1986). By contrast ，these leaves tions tions available to us ，we have only had an unclear demonstrate similar foliar characters to those of Alnus photograph photograph and simple description published by Zhang prenepalensis Hu & Chaney. As far as the so-called (1978) (1978) to go on. A clearer understanding of the F agus species from the locality third is concemed ，it structure structure of the fossil cupule would be very valuable is difficult to discuss it further because the original in in the interpretation of the systematic relationships of authors did not provide a description ， or any plates 174 174 植物研究雑誌第71 巻第3 号 平成 8 年 6 月 and figures ，and their fossilleaf was not available for Fushun taxon ，B etula populoides Hollick ， with regard exammat lO n. to its leaf base and the course of the secondary veins In In any case ， the name F agus chinensis Li is a later at the leafbase. is It ，however ，difficult to confirm this homonym ，having already been used by Oliver in identification ， because the fossil has no leaf apex. Hu 1890 1890 (Chang and Huang 1988) for an extant species. and Chaney (1940) suggested the specimen resem- Although this species has since been reduced to syn- bled Tilia miohenryan αHu & Chaney from the early onymy under F. longipetiolat αSeem.， ，one is not middle Miocene Shanwang f1 ora. permitted permitted to use F. chinensis for the fossil under the Material: Depository unclea r. Intemational Intemational Code of Botanical Nomenclature. Locality and age: Fushun， Liaoning Province ， From the f1 0ristic point of view ， the Eocene f1 0ra northeastem China ， Middle-Late Eocene. of Fushun Coalmine was considered to resemble the living living vegetation of eastem Sichuan (southwestem The Geological History of Chinese F agus China) and westem Hubei (central China) provinces It is often rather difficult to assess evolutionary (WGCPC 1978). In these areas ，F agus presently forms trends for given taxa because of the following factors a dense mixed forest with Betula ，Carpinus ，Acer ， though plenty of fossils may be available: Populus ，Platyc αry α，Alnus ，Li thocarpus ，Quercus First ， convergence of angiospermous mo 中hologi- and Castanopsis ，etc. These genera can also be found cal features and differing rates of evolutionary change fossil fossil in the Fushun f1 ora. This means that one might in the various organs (e.g.leaves ，woods ，fruits ，seeds ， expect to find Fagus in the fossillocality. However pollen and spores ，etc.) often lead to strong mosaic neither neither fossil pollen nor megafossils of F agus have evolution of plants and give rise to complicated ever been recovered. This could suggest that plant phylogenies. Consequently ，schemes based on a sin- components associated with F agus have changed in gle organ are less likely to re f1 ect the true phylogeny the the course of geological time. than those founded on a number of organs (see Material: Material: Housed at the Institute ofBotany (Beijing) Upchurch and Dilcher 1990). Unfortunately ，fossil and Nanjing Institute ofGeology and Palaeontology ， plants are usually preserved as isolated organs ， such Academia Sinica. as leaves ，pollen ，fruits and so on. This is the case in Locality Locality and age: Fushun ，Liaoning Province ， all the Chinese F agus documented so fa r. As the northeastem China ，Middle-Late Eocene (Guchengzi connections between the individual organs are con- Formation); Formation); Shanwang ，Shandong Province ，eastem jectural ， the resulting interpretation can be very one- China ， early middle Miocene (Shanwang Formation); sided. Tengchong ，Yunnan Province ，southwestem China ， Secondly ， plant taphonomic work suggests that late late Miocene to early Pliocene. not all plants/plant parts are buried and become fossil- 2) 2) Dicotylophyllum sp. ized (Ferguson 1985 ， Spicer 1991). This leads to a Fagus ?feroniae Ung. ，Florin ，Svensk Bo t. Tidsk. 12 ， further loss of information. l. pl. 1 ，figs. 5-7 (1 922). Finally ，it is dependent on the degree of sampling Discussion: Discussion: Fossil leaves described by Florin and study to which the fossil assemblages have been (1 920) as Dico η lophyllum from the Fushun Coal- submitted. To some extent ，discoveries of fossil plants mine are very different from F agus in leaf shape ， are accidenta l. Moreover paleobotanists are not infal- margin ，and venation. One of the specimens (Fl orin lible. Dilcher (1971) concluded that sixty percent of 1922 ， p l. 1， fig. 7) seems to be similar to another the taxa in the Eocene Wilcox f1 0ra of the southeastem June June 1996 Joumal of Japanese Botany Vo l. 71 No. 3 175

Table Table 2. Geological history of Fagus megafossils in China (E = early; M = middle; L = late; Qp = Pleistocene; Qh Holocene) = Holocene)

Time Time Eocene Oligocene Miocene Pliocene Quatemary

E M L E L E M L E L Qp Qh

F. F. sp. 1 F.g αlbanifolia F. F. sp. 2 F. F. stuxbergi F. F. engleriana F. F. praelucida F. F. sp. 3

U.S.A. U.S.A. were misidentified by Berry (1 916 ， 1930). that the world-wide earliest reliable record of Fagus Another Another good example is the Oligocene Creede flora pollen (F α gus granulata Piel) is from the lower from southem Colorado ， U.S.A. studied by Axelrod Oligocene of Canada. Recent work by Walther and (1 987) and Wolfe and Schom (1 990) respectively. Zetter (1993) shows that the oldest of record F agus in While Axelrod (1987) recognized 19 species belong- Europe is from the lower Oligocene. ing ing to 6 genera ， or 2 families of conifers ， and 53 Of the Chinese megafossils ， five species display species species assigned to 37 genera ， or 21 families of similarities to F agus engleriana Seem. ， while the Dicotyledones ，Wolfe and Schom (1 990) had only 8 other two probably resemble another Chinese en- species species attributed to 4 genera ， or 2 families of conifers demic beech ， F. lucidl αRehd. & Wils. This perhaps

and 24 species belonging to 18 genera ， or 7 families of indicates that the “F agus engleriana' にlike leaves are Dicotyledones. Dicotyledones. Thus different specific concepts common during the Cenozoic in China. Moreover ， splitters" (“ splitters" versus “lumpers ウalso cloud the picture. these fossils demonstrate low diversity of beeches in Because of the limitations mentioned above ，this the geological past in China. This is quite different account account of F agus based on individual fossils might be from that of the recent case. More materials ，particu-

changed changed when new material is recovered. As dis- larly with anatomically preserved specimens ， are cer 聞 cussed cussed above ， seven species ofbeech megafossil have tainly needed to further understand the early evolu- been been documented in China. Their geological distribu- tion of Fagus. tion tion is shown in Table 2. According According to the information presented in Table 2， We thank Profs. Shuang-Xing Guo (Na 吋ing ， the the current data on the fossil records of F agus ぽ e China) ，Ge Sun (Nanjing ， China) and Ji-Hui Zhang insufficient insufficient to reveal the clear history of the genus in (Guizhou ， China) for providing some photos and China. China. However ，F agus sp. 1 from the Late Eocene to related literature. Special thanks are also due to Prof. Early Early Oligocene seems to represent the oldest David K. Ferguson and D r. Reinhard Zetter (Vienna ， megafossil megafossil record 田 nong Chinese beech megafossils. Austria) for their comments and discussions. This F agus fossil reported previously from East Asian and work is mainly supported by the J apan Society for the westem sides of the Pacific basin were dated back to Promotion of Science Postdoctoral Fellowship (no. the the Oligocene (Tanai 1974 ， see also Taylor 1990). 94090) (YSL) ， the Ministry of Education ， Science Mu l1 er (1981) mentioned ， based on fossil pollen data ， and Culture of Japan no. 94090 (AM) and par t1 y 176 176 植物研究雑誌第71 巻第3 号 平成 8 年 6 月 financed from the Director Foundation of Nanjing 204-217. Liu Liu Y. S. in press a. Foliar architecture of Betulaceae and a Institute Institute of Geology and Palaeontology ，Academia revision revision of Chinese betulaceous fossil leaves. Palaeonto- Sinica ，which are greatly acknowledged. graphica Ab t. B. 一一一一一 in press b. Neogene floras ofChina. In:: Li X. X. et al.

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劉裕生，百原新，梅盛臭:中国産ブナ属 (ブナ科)大型化石の再検討 中国からこれまで報告されたブナ属大型化石を 界で最古のブナ属化石記録であろう. ブナ属とし 再検討したところ， 7 種類が確認された.このう て記載された F. chinesis Li と F. ? feroniae Unger ち， 1種類は殻斗の化石で，他の 6 種類は葉の化 は， ブナ属とは異なることが今回明らかになった. 石である . Fagus sp. 1は現世種の中国固有種 F. 新生代を通して“F. engleriana" タイプの葉化石 engleriana engleriana Seem. と F. longipetiolata Seem. に類似 が中国では普通に産出する.現在の中国と比べて， し，貴州省の後期始新世から前期漸新世の地層か ブナ属化石の多様性は乏しかったといえる. ら報告されている. これは，中国だけではなく世