vegetational history of the korean peninsula

© 2000 Blackwell Science Ltd. http://www.blackwell-science.com/geb 391

RESEARCH ARTICLE

Global Ecology & Biogeography (2000) 9, 391–402

Blackwell Science, Ltd

Vegetational history of the Korean PeninsulaWOO-SEOK KONG Department of Geography, KyungHee University, Seoul, 130–701, Korea (ROK). E-mail: [email protected]

ABSTRACT

1 The vegetational and environmental historyof the Korean Peninsula has been reconstructedby the use of both macrofossils and pollendata. These data were analysed within the time-frame between the Permian and the presentday.2 The continuous appearances of presentlyoccurring conifers and dicotyledons since theCretaceous indicate both the absence of cata-strophic environmental changes in the past, anda degree of long-term climatic continuity. Therewere, however, climatic fluctuations occurringwithin the full time-frame considered.3 The presence of a relatively rich flora in both

the Oligocene and Miocene suggests the exist-ence of a warm climate. The removal of theTertiary and Quaternary conifers from theHolocene deposits and the continued survival ofcryophilous conifers since the later Pleistocenemay be due to climatic deterioration.4 Overall, an analysis on the vegetational his-tory seems to provide information for the betterunderstanding of the present vegetation as wellas providing data on environmental change inthe region.

Key words Climatic fluctuations, conifers, dico-tyledons, Korean Peninsula, macrofossils, pollen,Quaternary period, Tertiary period, vegetationaland environmental history.

INTRODUCTION

This present work aims to reconstruct thevegetational and environmental history of Koreaand also to analyse the formation process ofpresent-day assemblages of Korean conifers anddicotyledons. To understand the vegetationalhistory, both macrofossil and pollen data fromthe stratigraphy of both the ROK (Republic ofKorea; the official name for South Korea) andthe DPRK (Democratic People’s Republic ofKorea; the official name for North Korea) havebeen collected. Overall, 40 sites for the conifersand 75 sites for the dicotyledons (Fig. 1) havebeen selected (Nan, 1984; Rim et al., 1994; Kong,1995, 1996, 1997). Macrofossils for the Pre-UpperMiocene and pollen data since the UpperMiocene have been analysed within the timeframe between the Permian and the present day.The selected data are discussed on the basis ofstratigraphic chronology in respect of the pre-Holocene deposits as well as with respect to

absolute ages for those Pleistocene and Holocenedeposits for which radiocarbon-dating data areavailable.

VEGETATION OF THE PALAEOZOIC ERA

The appearance of the oldest conifers in Koreawas in deposits recovered at Sadong. These dateback to the Permian period, and include Elato-cladus, Ullmannia and Walchia (Lee, 1987). How-ever, only Elatocladus reappeared at Kobangsanduring the Triassic period (Fig. 1).

VEGETATION OF THE MESOZOIC ERA

Out of 13 conifers from the Mesozoic era(Lee, 1987), Triassic and Jurassic conifers includeAraucarites at Kangso (DPRK), Palissya at Kimpo(ROK), Pityophyllum at Kimpo, Youngwol, Munk-yong (ROK) and Pyongyang (DPRK), Stenorachis

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© 2000 Blackwell Science Ltd, Global Ecology & Biogeography, 9, 391–402

Fig. 1 Locations of fossil plants discovered on the Korean Peninsula.


Vegetational history of Korea 393


at Nampo (DPRK), Schizolepis at Youngwol,and Swedenborgia at Youngwol and Pyongyang.Lower Cretaceous conifers consist of Brachyphyllumat Taegu (ROK), as well as Sariwon and Cheryong(DPRK), Pityophyllum at Chinju (ROK) andTaegu, Cyparissidium at Chinju, Czekanowskiaat Chinju, and Sequoia and Xenoxylon at Taegu.Conifers of the Upper Cretaceous are Freno-lepsis at Youngdong, Sansudong of Chinan,Taegu, Sariwon and Cheryong, Brachyphyllum atYoungdong, Sansudong of Chinan, Sariwonand Cheryong, and Pinus at Dalgil of Chinan,Kyongsang (ROK), Youngdong, Sariwon andCheryong. However, only Pinus has thrived asone of the dominant trees since the Cretaceous,along with Ginkgo (Kong, 1995).

The oldest dicotyledons on the Korean Pen-insula date back to the Cretaceous and include16 genera: Platanus, Viburnum, Populophyllum,Tabeinidium at Dalgil and Sansudong of Chinanand Phyllites, Zamiophyllum at Hwasoon, as wellas Salix, Populus, Nelumbies, Menispermites,Cinnamomum, Ilex, Rhamnites, Grewia, Aralia andLindera at Youngduk (ROK). Nine genera, Platanus,Viburnum, Salix, Populus, Cinnamomum, Ilex,Grewia, Aralia and Lindera, still grow naturallyin the wild (Kong, 1996).

VEGETATION OF THE CENOZOIC ERA

Oligocene vegetation

No deposits with plant fossil data are availablefor the Palaeocene and Eocene, and neither haveany conifers been reported from the Oligocenein Korea. However, some indications as to whattypes of vegetation were present are availablefrom NE Asian material collected over a some-what broader range of territory. In these epochs,due to the existence of generally warm climates, anevergreen broad-leaved vegetation has dominated,extending to latitude 60°N during the Eocene(Wolfe, 1980). According to Tanai (1972), manyother present-day Korean genera were alreadypresent in East Asia during the Upper Eocene,among which were Platcarya, Castanopsis, Quercus,Ficus, Machilus, Pittosperum, Ilex, Zizyphus,Ternstroemia, Xylosma and Ligustrum.

Twelve dicotyledonous taxa from Oligocenedeposits at Pongsan (Rim et al., 1994) include

Populus (three spp.), Myrica, Juglans, Ficus,Credneria, Platanus (three spp.), Celastrus, Acer,Sapindus, Zizyphus, Hedera and Viburnum (twospp.). The appearance of thermophilous or warmth-loving evergreen broad-leaved trees during theOligocene at c. 38°30′N, along with a comparisonof the present-day northern limits of these, namelyMyrica at 34°30′N, Ficus at 35°30′N and Hederaat 37°30′N (Kong, 1992), is indicative of the factthat the Oligocene climate was somewhat warmerthan the climate of today.

Miocene vegetation

For conifers of the DPRK, out of 15 genera ofthe Lower Miocene at Yongdong (Rim et al.,1994), three genera (Picea, Glytostrobus andMetasequoia) are recognized. For dicotyledons,out of 15 trees in toto, the Lower Miocene floraat Yongdong consists of 11 genera: Pterocarya,Betula, Carpinus, Ostrya, Zelkova, Cercidiphyllum,Rosa, Acer (four spp.), Aesculus, Tilia (four spp.)and Alangium.

In the ROK, out of 44 trees of the LowerMiocene flora (Huzioka, 1943, 1951, 1972;Bong, 1979; Chun, 1996) at Changgi, five con-ifers, namely Picea, Pseudotsuga, Glyptostrobus,Metasequoia and Sciadopitys, have been reported.Sixteen conifers, Keteleeria, Pinus, Picea, Abies,Larix, Psudolarix, Cedrus, Tsuga, Araucaria,Cryptomeria, Sciadopitys, Sequoia, Taxodium,Cupressus, Libocedrus and Juniperus, are alsonoted out of 44 genera at Kumkwangdong,near Changgi (Bong, 1979). Seventeen conifergenera were recognized out of 61 genera of theLower Miocene at Kampo-Eoil (Bong, 1980),namely Pinus, Picea, Abies, Larix, Cedrus, Tsuga,Araucaria, Cephalotaxus, Taxus, Podocarpus,Cryptomeria, Sciadopitys, Sequoia, Taxodium,Cupressus, Libocedrus and Juniperus.

Thirty-nine dicotyledons from 44 genera havebeen discovered at Changgi of the Lower Miocene.These include Alnus (two spp.), Betula (three spp.),Carpinus (four spp.), Corylus, Castanea, Fagus(two spp.), Nothofagus (two spp.), Quercus (fourspp.), Populus, Salix (two spp.), Carya (twospp.), Juglans (three spp.), Liriodendron, Morus,Ulmus, Zelkova, Tilia (two spp.), Ilex (two spp.),Acer, Fraxinus, Symplocarpus, Platycarya, Ostrya,Hydrangea, Magnolia (two spp.), Sorbus (twospp.), Cladrastis, Rhus, Acer (five spp.), Aesculus,

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Alangium, Hemitrapa, Cornus, Rhododendron, Dio-spyros, Viburnum, Ficus, Rhamnus and Liquidambar.The Changgi flora contains 33 trees in com-mon with the known Miocene flora of Japan, butonly seven trees are in common with those ofthe Shantung Province of China (Huzioka, 1951,1972), immediately to the west, across the YellowSea. Another Lower Miocene flora from Kampo-Eoil comprises 23 dicotyledons from 61 treegenera in toto, especially Alnus, Betula, Carpinus,Ostrya, Corylus, Castanea, Fagus, Nothofagus,Quercus, Populus, Carya, Juglans, Pterocarya,Liriodendron, Morus, Ulmus, Liquidambar, Tilia,Ilex, Acer, Fraxinus, Sapindus and Cinnamomum.

In the DPRK at Tongchon (Rim et al., 1994),three conifers from 15 genera of the MiddleMiocene are Pinus, Glytostorobus and Metasequoia.Seven conifers of the same period from Hoeryong,among 31 genera overall, include Picea, Pinus,Sequoia, Taxodium, Glyptostrobus, Metasequoiaand Sciadopitys. Other Middle Miocene conifersat Kokonwon comprise seven genera, Keteleeria,Pseudolarix, Glytostrobus, Metasequoia, Sciadopitys,Calocedrus and Thujopsis, from 30 genera. Threeconifers (Picea, Glytostrobus and Metasequoia)out of 30 genera of the same stage have been dis-covered at Hamjindong.

Middle Miocene dicotyledons at Tongchonconsist of 12 genera, Populus, Salix (two spp.),Comptonia, Pterocarya, Alnus, Betula, Fagus,Zelkova, Cercidiphyllum, Acer (five spp.), Hemi-trapa and Alangium, among 15 tree genera.Seven Middle Miocene Yoosun dicotyledonsfrom 31 genera contain Carpinus, Cornus, Acer,Populus, Platanus, Fagus and Myrica. AnotherMiddle Miocene deposit at Hangyoung includes19 dicotyledons: Castanea, Fagus (three spp.),Quercus (six spp.), Engelhardtia, Pterocarya, Betula,Carpinus (three spp.), Alnus, Planera, Ulmus,Zelkova, Cercidiphyllum, Platanus (two spp.),Hamamelis, Ailanthus, Cedrela, Populus, Acer(three spp.) and Alangium.

In Kokonwon from Middle Miocene deposits,21 dicotyledons from 30 genera have been dis-covered: Salix, Engelhardtia, Alnus, Betula (twospp.), Carpinus, Castanea, Fagus (four spp.), Quercus(three spp.), Cyclobalanopsis, Zelkova, Cercidiphyllum,Liriodendron, Platanus, Gleditisia, Ilex, Euonymus,Acer (four spp.), Vitis, Kalopanax, Porana andAlangium. The Hamjindong flora of the MiddleMiocene contains 24 dicotyledons: Populus,

Juglans, Pterocarya, Alnus, Carpinus (two spp.),Fagus (three spp.), Ulmus (two spp.), Zelkova,Cercidiphyllum, Liriodendron, Lindera, Sassafras,Platanus, Hamamelis, Parrotia, Prunus, Cercis,Acer, Aesculus, Tilia, Rhododendron, Diospyros,Styrax and Fraxinus. From 30 genera in theChangpungni deposit of the same period arefound 12 dicotyledons: Carpinus, Fagus, Compto-nia, Salix, Juglans, Cinnamomun, Acer, Corylus,Viburnum, Vitis, Prunus and Rhamnus.

In the ROK, Middle Miocene deposits atYonil contain 18 conifers from 60 genera of theYonil flora (Bong, 1982): Pinus, Picea, Abies,Larix, Pseudolarix, Cedrus, Keteleeria, Tsuga,Pseudotsuga, Araucaria, Sciadopitys, Sequoia,Taxodium, Podocarpus, Dacrydium, Cupressus,Libocedrus and Juniperus.

Also in the ROK, 11 conifers from 21 generaof the Upper Miocene at Pukpyong (Yu, 1971)include Abies, Larix, Tsuga, Picea, Pinus, Meta-sequoia, Cedrus, Pseularix, Keteleeria, Podocarpusand Taxus.

During the Upper Miocene, 39 dicotyledonshave been reported at Yonil from the 60 treegenera. These were Alnus, Betula, Carpinus (threespp.), Castanea, Fagus, Nothofagus, Quercus,Populus, Carya, Juglans, Magnolia, Ulmus, Tilia,Erica, Myrica, Liquidambar, Ilex, Acer (three spp.),Fraxinus, Cornus, Comarum, Pterocarya, Castano-psis, Cyclobalanopsis (three spp.), Pasania (twospp.), Zelkova, Cinnamomum, Cryptocarya, Lindera,Phoebe, Platanus, Parrotia, Entada, Sapindus,Paliurus, Alangium, Hemitrapa, Rhododendronand Carpites. In Pukpyong of the same period,there are 14 dicotyledons, namely Carpinus,Quercus, Betula, Fagus (two spp.), Pasania, Alang-ium, Styrax, Rhus, Rhododendron, Salix, Ilex,Trochodendron, Tilia and Platycarya.

Overall, 10 conifer genera of the KoreanPeninsula survived from the Miocene but becameextinct at the end of the Tertiary. These wereGlyptostrobus, Keteleeria, Calocedrus, Thujopsis,Araucaria, Libocedrus, Pseudolarix, Cedrus,Podocarpus and Dacrydium. Conifers that flour-ished from the Miocene to the Pleistocene, butare now extinct, include Pseudotsuga, Metasequoia,Sciadopitys, Taxodium, Cryptomeria and Cupres-sus (Kong, 1995).

The Miocene dicotyledons of Korea contain72 genera: Platanus, Viburnum, Salix, Populus,Cinnamomum, Ilex, Lindera, Myrica, Juglans,




Ficus, Acer, Sapindus, Carpinus, Cornus, Fagus,Castanea, Quercus, Engelhardtia, Pterocarya, Betula,Alnus, Planera, Ulmus, Zelkova, Cercidiphyllum,Hamamelis, Ailanthus, Cedrela, Alangium, Cyclo-balanopsis, Liriodendron, Gleditsia, Euonymus,Vitis, Kalopanax, Porana, Ostrya, Rosa, Aesculus,Tilia, Sassafras, Parrotia, Prunus, Cercis, Rhododen-dron, Diospyros, Styrax, Fraxinus, Comptonia,Hemitrapa, Corylus, Nothofagus, Carya, Morus,Platycarya, Hydrangea, Sorbus, Cladrastis, Rhus,Rhamnus, Liquidambar, Erica, Magnolia, Castano-

psis, Paliurus, Pasania, Cryptocarya, Phoebe,Entada, Carpites and Trochodendron (Kong,1996, 1997). Out of those, 53 genera still growin Korea. Survey data for a selection of generaof phytogeographical importance are presentedin Table 1.

The presence of a relatively rich flora duringthe Miocene (Table 1), including thermophilousevergreen broad-leaved trees, namely Castanopsis,Cyclobalanopsis, Cinnamomum, Ilex and Myrica,suggests the continuation of a warm climate in

Table 1 Presence of selected fossil conifers and dicotyledons on the Korean Peninsula

GeneraPale.Per.

Mesozoic Cenozoic

Tri. Jur. Cre. Pal. Eoc. Oli. Mio. Ple. Hol. Pre.

Walchia* ×Sequoia* × Pinus (Diplo.) × × × × ×Metasequoia* × ×Picea × × × ×Abies × × × ×Tsuga × × ×Juniperus × × × ×Larix × × ×Cephalotaxus × ×Taxus × × ×Thuja × ×Pinus (Haplo.) × ×Salix × × × × ×Cinnamomum × × × ×Ilex × × × × ×Juglans × × × × ×Acer × × × × ×Carpinus × × × ×Quercus × × × ×Betula × × × ×Alnus × × × ×Tilia × × × ×Rhododendron × ×Fraxinus × × × ×Sorbus × × ×Magnolia × × ×Celtis × × ×Benzoin × ×Buxus × ×Daphne × ×Acanthopanax × ×

* Extinct genera in Korea. Kong (1994, 1995, 1996). Abbreviations: Pale. = Palaeozoic, Per. = Permian,Tri. = Triassic, Jur. = Jurassic, Cre. = Cretaceous, Pal. = Palaeocene, Eoc. = Eocene, Oli. = Oligocene,Mio. = Miocene, Ple. = Pleistocene, Hol. = Holocene, Pre. = Present, Diplo. = Diploxylon, Haplo. = Haploxylon.


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some parts of the Korean Peninsula since then(Kong, 1996). Unfortunately, however, from thePliocene to the Upper Pleistocene, no depositswith fossil plants are available from Korea, andso a hiatus in the vegetation data exists of someconsiderable length.

Pleistocene and Holocene vegetation

In the DPRK (Rim et al., 1994), conifers of theLower Pleistocene at Hwasung, Eorang, Saepoand Hyoiyang comprise Pinus, Tsuga and Scia-dopitys. Those of Haesang of the same period(Kim, 1992) consist of 31 taxa, and 24.4% of thepollen sum are conifers, including Cupressaceae(12.4%), Pinus (9.3%), Taxodiaceae (1.6%) andLarix (1.0%). Dicotyledons of the Lower Pleis-tocene at Hwasung consist of 15 genera: Myrica,Pterocarya, Magnolia, Fagus, Castanea, Ilex,Juglans, Rhus, Betula (two spp.), Quercus, Populus(two spp.), Acer (two spp.), Alnus, Castanea andCarpinus. Out of 31 genera at Haesang of theLower Pleistocene, Betula and Quercus comprise55.6% of the pollen sum of dicotyledons, alongwith Ulmus, Tilia, Salix, Populus, Pterocarya,Carya, Engelhardtia, Alnus, Ostrya, Celtis, Liqui-dambar, Ilex, Elaeagnus and Corylus.

In the DPRK too, the Middle Pleistoceneconifers at Saebyol include Pinus (10.4% of thepollen sum), Cupressaceae (5.2%), Picea (4.4%),Taxodiaceae and Tsuga. Dicotyledons of theMiddle Pleistocene at Saebyol include Juglans,Quercus, Castanea, Magnolia and Fagus.

Furthermore, at Yonggok in the DPRK ofthe Upper Pleistocene, out of 18 families or24 genera (Han, 1995), 32% of the pollen sumbelongs to conifers. These are as follows: Pinus(13.7%), Picea (4.4%), Abies (1.7%), Cryptomeria(3.7%), Larix (2.6%), Tsuga (1.2%), Metasequoia(1.2%), Pinaceae (1.9%), Taxodiaceae (0.6%) andCupressaceae (0.6%). Upper Pleistocene depositsat Seungnisan consist of 19 families or 15 generaof pollen. Conifers include Pinus (51.4% of thepollen sum), Cupressaceae (8.25%), Larix (3.6%),Abies (1.8%) and Picea (1.8%). Out of sevenfamilies or 16 genera of Hwadae pollen, 28.5%of the pollen sum belongs to conifers, Pinus(10.5%), Larix (5%), Picea (4%), Abies (1%),Juniperus (4%) and Thuja (1.5%). The presence ofa large number of conifers in the pollen record,and the dominance of those genera, including

cryophilous or cold-tolerant ones, indicate thatthe climate then was somewhat cooler than theclimate of today.

Out of 18 families or 24 genera in the Yonggokflora of the Upper Pleistocene, 16 are dicoty-ledons: Magnolia, Ulmus, Zelkova, Fagus, Quercus,Betula, Alnus, Carpinus, Juglans, Pterocarya,Carya, Ilex, Myrica, Tilia, Acer and Liquidambar.Out of 19 families or 15 genera of the UpperPleistocene deposit at Seungnisan, dicotyledonsconsist of seven genera, Quercus, Betula, Tilia,Salix, Juglans, Ulmus and Corylus. From sevenfamilies or 16 genera in Upper Pleistocenedeposits at Hwadae are reported Salix, Betula,Alnus and Quercus.

In the ROK, the Upper Pleistocene depositat Durubong (Park, 1984) contains pollen of 13families or 12 genera. Out of them, 23.9% ofthe pollen sum are conifers of the genera Taxus,Abies, Larix and Pinus. From 22 families or 24genera at Chummal-Yonggul, conifers consist ofTaxus, Abies, Pseudotsuga, Tsuga, Picea, Larix,Pinus and Cupressaceae. According to Youn(1996), Picea was the predominating genus sincec. 57 000 years at Youngyang and a relativelyhigh proportion of conifers are also noticedfor Pinus, Abies and Larix. The flora of Kajofrom c. 32 050–30 700 years (Jo et al., 1987)consists of 88 pollen types, including conifersof the genera Picea, Pinus, Abies and Larix.From 10 families or six tree genera, two conifers(Abies and Pinus) are noted at Sokjangni fromc. 29 000 years . From c. 17 000–15 000 years

at Younglangho (Kim, 1980), Picea, Larix,Abies and Pinus (Haploxylon and Diploxylon)form about 10% of the total pollen sum.

Out of 22 families or 24 genera at Chummal-Yonggul (Upper Pleistocene) 19 dicotyledonpollen types were recorded: Salix, Juglans, Betula,Castanea, Fagus, Quercus, Celtis, Benzoin, Ilex,Tilia, Smilax, Populus, Alnus, Carpinus, Ulmus,Cinnamomun, Prunus, Elaeagnus and Aralia. AtDurubong for the Upper Pleistocene, 13 familiesor 12 genera were recorded, including seven dicoty-ledons, namely Alnus, Betula, Carpinus, Quercus,Acer, Tilia and Ligustrum. From c. 43 000–18 000years at Youngyang, eight dicotyledons, Quer-cus, Castanea, Ulmus, Zelkova, Corylus, Carpinus,Tilia and Acer, occurred.

Out of 88 genera at Kajo, c. 30 000 years, dicotyledons include 32 genera: Akebia,




Stauntonia, Juglans, Myrica, Alnus, Betula,Corylus, Castanea, Castanopsis, Quercus, Ulmus,Hydrangea, Corylopsis, Pittosperum, Rosa, Raphi-dephis, Phellodendron, Poncirus, Sapinium, Buxus,Daphne, Rhus, Acer, Euonymus, Tilia, Salix, Pun-ica, Acanthopanax, Symplocos, Campsis, Callicarpaand Vitex. From c. 32 000, Fraxinus occurred atKumkul, and out of 10 families or six generain the pollen record c. 28 000–25 000 years atSokjangni Alnus, Betula, Magnolia, Acer andLigustrum occurred. The dicotyledons of Suyanggaeof c. 25 000–18 000 years , and of Changnaeof the same period, include Salix, Alnus, Car-pinus, Betula, Quercus, Castanea, Ulmus, Celtisand Ligustrum; and Quercus, Prunus and Salix,respectively.

According to Grichuk (1984), vegetation inthe Far East of Russia during the Last GlacialMaximum, displayed regional variations. It con-tained montane subarctic tundra, subalpine andalpine meadows, open Betula and Picea forests,a light coniferous montane forest, a dark conif-erous and Betula montane forest and conifer-ous and broad-leaved montane forest.

In NE China, the southern limit of theboreal forest moved southward to extend fromthe southern part of the Liadong Peninsula tothe mountain region of northern Hebei. Inturn, to the south of this line, a coniferous anddeciduous temperate forest, or steppe forest,extended southward to the Yangtze Delta. Dur-ing the late Pleistocene, the climate was coolerand drier than today (Zhang, 1984; Jiang &Piperno, 1999).

In Japan, tundra vegetation occupied thearea to the north of 44°30′N and was alsopresent in the south above 900 m a.s.l. The restof the area was covered generally with morespecific boreal or subalpine plants, domin-ated by Pinus pumila and Betula ermanii. How-ever, in climatically more favourable locations,Picea, Betula ermanii, Pinus, Larix and otherboreal and bog taxa were common features ofthe pollen data (Yamanaka, 1978; Igarashi &Kumano, 1981; Tsukada, 1983a). Northern andcentral Honshu contained a subalpine coniferousforest, mainly comprised of Picea, Abies andTsuga (Quaternary Flora Research Group, 1974;Yasuda, 1978; Sakai, 1981; Tsukada, 1983b).

During the Holocene, on the eastern partof the ROK (Kim, 1980), from c. 10 000–6700

years , Pinus (Diploxylon) and Abies occurredat Younglangho, along with Pinus (Diploxylon),which accounts for up to 50% of the totalpollen sum. It became a dominant tree alongwith Pinus (Haploxylon) and Abies during theperiod of c. 6700–4500 years . The declineof Pinus (Diploxylon) pollen from 50% to 10%since c. 1400 years is noticeable, but Abiesand Pinus (Haploxylon) show no drastic changes.The appearance of charcoal and cultivated ricepollen in the deposits implies the presence ofanthropogenic disturbance of the natural vegeta-tion (Kong, 1994; Park et al., 1996).

In Chumunjin, Pohang, Bangojin and Yeian(Jo, 1987) the dominance of Pinus, however,continued at up to 90% of the pollen sum fromc. 7800 down to 770 years , along with thepresence of other conifers, such as Abies andPicea. The increase of Pinus pollen is also foundin the western part of the ROK. For example,5–15% of the pollen sum at Hwangdung fromc. 6260–4950 years , at Ilsan from c. 5650–2270 years , at Muan 45–90% of the pollensum, and at Pyongtaek, Chollipo, Kimje andMiruksaji. The admixture of Pinus, Abies andPicea is also noticed from c. 2200 years at theDaeamsan high moor (1316 m a.s.l.) in centralKorea (Kang, 1980).

The proportion of dicotyledons at Young-langho (Kim, 1980) in east Korea rapidlyincreased in this period, for example to over20% of the pollen sum for Quercus. Other dicoty-ledonous genera include Carpinus, Salix, Ulmus,Celtis, Juglans, Carya, Tilia, Alnus, Castaneaand Fagus at approximately 10 000–6700 years. Twenty-five per cent of the pollen sum wasQuercus, along with Carpinus, Ulmus, Celtis, Alnusand Juglans at c. 6700–4500 years . Thirty-fiveper cent of the pollen sum was Quercus, alongwith Carpinus, Corylus, Alnus, Ulmus, Betulaand Juglans at c. 4500–1400 years . Forty percent of the pollen sum was Quercus along withAlnus, Betula, Corylus, Carpinus and Juglansat approximately 1400 years . Data from Pohangfrom c. 9800 years (Jo, 1979; 1987) consistof 13 dicotyledons: Salix, Juglans, Carya, Alnus,Betula, Carpinus, Corylus, Castanea, Fagus,Quercus, Ulmus, Zelkova and Tilia. The per-centage of Quercus increased from 20% of thepollen sum at c. 9820 to 40% of the pollensum at c. 7810 years .


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Pollen assemblages of dicotyledons in otherareas of the ROK vary from site to site. From6270 to 4950 years , eight genera (Juglans,Platycarya, Alnus, Carpinus, Corylus, Castanea,Fagus and Quercus) were present at Hwangdung.At c. 6000–2600 years , 11 genera (Quercus,Alnus, Fraxinus, Carpinus, Juglans, Castanea,Ulmus, Corylus, Salix, Betula and Tilia) havebeen reported from Ilsan. At c. 4060–770 years, 18 genera (Quercus, Alnus, Corylus, Castanea,Carpinus, Ulmus, Myrica, Tilia, Salix, Betula,Cyclobalanopsis, Zelkova, Juglans, Platycarya,Fraxinus, Acer, Castanopsis and Celtis) haveoccurred at Bangojin. From c. 4000 years , 17genera (Lespidobalanus, Carpinus, Cyclobalanopsis,Castanopsis, Celtis, Corylus, Fraxinus, Zelkova,Ulmus, Betula, Acer, Juglans, Myrica, Platycarya,Tilia, Elaeagnus and Salix) have been discoveredat Wolhamji.

From c. 3000 years , 15 genera (Ligustrum,Quercus, Ulmus, Alnus, Carpinus, Corylus, Betula,Tilia, Juglans, Viburnum, Fraxinus, Fagus, Ilex,Salix and Castanea) have occurred at Pyongtaek.From c. 2200 years , nine genera (Quercus,Betula, Alnus, Carpinus, Corylus, Tilia, Salix,Juglans and Ulmus) were reported at Daeamsan.From c. 2000 years , nine genera (Salix, Juglans,Alnus, Betula, Carpinus, Corylus, Castanea, Fagusand Quercus) were present at Chumunjin. For thesame period, 18 genera (Platycarya, Juglans, Alnus,Betula, Carpinus, Corylus, Castanea, Castanopsis,Quercus, Cyclobalanopsis, Ulmus, Zelkova, Morus,Akebia, Rosa, Rhus, Tilia and Fraxinus) havebeen reported for Chollipo.

From c. 1500 years 11 genera (Quercus,Carpinus, Corylus, Cyclobalanopsis, Ulmus, Alnus,Castanopsis, Celtis, Zelkova and Myrica) havebeen discovered at Muan. From c. 1280 years, 13 genera (Quercus, Alnus, Carpinus, Ulmus,Elaeagnus, Vitis, Cyclobalanopsis, Salix, Juglans,Corylus, Zelkova, Celtis and Platycarya) havebeen reported at Yeian. Ten genera (Quercus, Alnus,Carpinus, Corylus, Acer, Castanopsis, Juglans,Betula, Ulmus and Platycarya) were present atKimje and nine genera (Quercus, Ulmus, Zelkova,Salix, Carpinus, Alnus, Corylus, Betula andCastanea) have been discovered at Miruksaji forthe same period.

Overall, of the conifers in Korea, Picea, Abiesand Juniperus appeared continuously from theMiocene to the Pleistocene and Holocene. Tsuga,

Larix and Taxus occurred from the Miocene tothe Pleistocene but not in the Holocene. Cepha-lotaxus is reported only from the Tertiary, andThuja and Pinus (Haploxylon) only from thePleistocene. Conifers that survived from theTertiary to the Quaternary, but which do notappear in the Holocene, include Sciadopitys,Taxodiaceae, Metasequoia and Cryptomeria.The extinction of those genera may be due tothe climatic deterioration during the laterPleistocene. Pinus, Juniperus, Abies, Picea andCupressaceae continued to exist from the Terti-ary to the later Pleistocene. The expansion ofPicea, Abies, Larix, Taxus and Thuja duringthe later Pleistocene appears to be influencedby the climatic cooling trends. Holocene con-ifers with phytogeographical importance includePinus (Diploxylon), Picea, Abies and Juniperus(Table 1).

During the Pleistocene, 54 dicotyledons,including pre-Pleistocene genera such as Salix,Populus, Cinnamomum, Ilex, Aralia, Myrica,Juglans, Acer, Carpinus, Fagus, Castanea, Quer-cus, Engelhardtia, Pterocarya, Betula, Alnus, Ulmus,Zelkova, Euonymus, Ostrya, Rosa, Tilia, Prunus,Fraxinus, Corylus, Carya, Hydrangea, Sorbus,Rhus, Liquidambar, Magnolia and Castanopsis, con-tinued to occur. There are also newly emergedPleistocene genera, such as Celtis, Elaeagnus,Benzoin, Smilax, Ligustrum, Akebia, Stauntonia,Corylopsis, Pittosporum, Raphidephis, Phelloden-dron, Poncirus, Sapium, Buxus, Daphne, Punica,Acanthopanax, Symplocos, Campsis, Callicarpa,Vitex and Lespidobalanus.

Twenty-eight dicotyledons of the Holoceneinclude Viburnum, Salix, Ilex, Myrica, Juglans,Acer, Carpinus, Fagus, Castanea, Quercus, Betula,Alnus, Ulmus, Zelkova, Cyclobalanopsis, Rosa,Tilia, Fraxinus, Corylus, Carya, Morus, Platycarya,Rhus, Castanopsis, Celtis, Eleagnus, Ligustrum andAkebia (Table 1).

The vegetational difference between the west-ern coast and eastern coast of Korea during theHolocene was noticeable. In a relatively gentlewestern Korea, Alnus-dominated vegetation con-tinued from c. 6250–1500 years , but Pinus tookover from Alnus subsequently. In a relativelymontane eastern Korea, however, Quercus andPinus dominated from c. 10 000–2000 years ,but Pinus became predominant from c. 2000 years. (Kong, 1994).




At present, six families, 16 genera and 66species of conifers, including 19 cultivated spe-cies of conifers, are reported in Korea. Theseare Cephalotaxus (two spp.), Torreya (one sp.),Taxus (three spp.), Pinus (21 spp.), Juniperus(13 spp.), Abies (four spp.), Cedrus (one sp.),Larix (four spp.), Picea (six spp.), Tsuga (onesp.), Chamaecyparis (three spp.), Thuja (one sp.),Cryptomeria (one sp.), Taxodium (one sp.) andMetasequoia (one sp.) (Lee, 1986). At present,the number of Korean dicotyledons reachesup to c. 2649 species (Park & Chung, 1996). Alarge number of conifers and dicotyledons atpresent on the Korean Peninsula may in part berelated to the long-term environmental stability,along with the periodic climatic fluctuations anddiverse habitats, which ensured the survival ofvarious conifers and dicotyledons.

DISCUSSION

Despite the general paucity of plant fossils overthe country as a whole, scattered data have allowedthe reconstruction of the vegetational and environ-mental history of the Korean Peninsula.

Pre-Cretaceous vegetation consists exclusivelyof conifers: Elatocladus, Ullmannia and Walchiafor the Permian period, and Araucarites, Palissya,Pityophyllum, Stenorachis, Schizolepis and Sweden-borgia for the Triassic and Jurassic periods.All of these genera, however, are now extinct

(Table 2). Out of 26 genera of the Cretaceousperiod, major or common vegetation componentsinclude Brachyphyllum, Sequoia, Pinus, Platanus,Viburnum and Populus, but 16 (62%) of them,Brachyphyllum, Pityophyllum, Cyparissidium,Czekanowskia, Sequoia, Xenoxylon, Frenolepsis,Ginkgo, Platanus, Populophyllum, Tabeinidium,Phyllites, Zamiophyllum, Nelumbies, Menispermitesand Rhamnites, are now extinct (Table 2).

The Oligocene period genera consist exclus-ively of dicotyledons, and major genera includePopulus, Myrica, Ficus, Platanus, Hedera andViburnum. Two genera (17%) of the Oligocenevegetation (Table 2), i.e. Credneria and Platanus,are now extinct.

The major genera of the Lower Mioceneperiod comprise Pinus, Picea, Metasequoia,Quercus, Fagus and Acer. Out of 65 conifers anddicotyledons, 21 genera (32%) are now extinct(Table 2), including 14 conifers: Glytostrobus,Metasequoia, Pseudotsuga, Keteleeria, Pseudolarix,Cedrus, Araucaria, Podocarpus, Cryptomeria,Sciadopitys, Sequoia, Taxodium, Cupressus andLibocedrus, and seven dicotyledons: Nothofagus,Liriodendron, Carya, Aesculus, Hemitrapa,Liquidambar and Pterocarya.

The vegetation of the Middle Miocene periodincludes 70 genera, including common generai.e. Pinus, Picea, Metasequoia, Fagus, Acer andCarpinus, but at present 31 of them (44%) areextinct (Table 2). Sixteen extinct conifers are

Table 2 Temporal change of vegetation on the Korean Peninsula

Eras Periods No. and ratio ofconifers

No. and ratio ofdicotyledons

Extinction ratio(no. of C : D)

Cenozoic Holocene 3 (9%) 30 (91%) 6% (0 : 2)

U. Pleistocene 14 (22%) 50 (78%) 16% (3 : 7)M. Pleistocene 5 (50%) 5 (50%) 0% (0 : 0)L. Pleistocene 6 (19%) 25 (81%) 16% (1 : 4)

U. Miocene 11 (19%) 46 (81%) 35% (5 : 15)M. Miocene 22 (31%) 48 (69%) 44% (16 : 15)L. Miocene 23 (35%) 42 (65%) 32% (14 : 7)

Oligocene 0 (0%) 12 (100%) 17% (0 : 2)

Mesozoic Cretaceous 10 (38%) 16 (62%) 62% (8 : 8)Tri. & Jura. 6 (100%) 0 (0%) 100% (6 : 0)

Palaeozoic Permian 3 (100%) 0 (0%) 100% (3 : 0)

Abbreviations: No. = number of genera, C = conifers, D = dicotyledons, U. = upper, M. = middle, L. = lower,Tri. = Triassic, Jura. = Jurassic.


400 W.-S. Kong


known: Glytostrobus, Metasequoia, Sequoia, Tax-odium, Sciadopitys, Cedrus, Keteleeria, Pseudot-suga, Araucaria, Podocarpus, Dacrydium, Cupressus,Libicedrus, Pseudolarix, Calocedrus and Thujopsis.Fifteen extinct dicotyledons comprise Comptonia,Pterocarya, Cercidiphyllum, Hemitrapa, Platanus,Engelhardtia, Hamamelis, Ailanthus, Cedrela,Liriodendron, Porana, Sassafras, Parrotia, Cercisand Aesculus.

The vegetation of the Upper Miocene periodconsists of 57 genera, and common generainclude Pinus, Abies, Metasequoia, Quercus, Fagusand Carpinus, but 20 genera (35%) are nowextinct. Extinct genera are comprised of: Meta-sequoia, Cedrus, Pseudolarix, Keteleeria, Podocar-pus, Nothofagus, Carya, Erica, Liquidambar,Comarum, Pterocarya, Pasania, Cryptocarya,Phoebe, Platanus, Parrotia, Entada, Hemitrapa,Carpites and Trochodendron.

The Lower Pleistocene period vegetation con-tains 31 taxa, and the major element includesPinus, Larix, Cupressaceae, Populus, Alnus andIlex, but five taxa (16%), Sciadopitys, Ptero-carya, Carya, Engelharditia and Liquidambar,are now extinct. Out of 10 Middle Pleistoceneperiod taxa, all still grow in the wild (Table 2),and common elements were Pinus, Cupressaceae,Picea, Juglans, Quercus and Castanea.

The vegetation of the Upper Pleistocene periodincludes 64 taxa (Table 2), and the major generaare comprised of: Pinus, Picea, Larix, Quer-cus, Betula and Alnus. Ten extinct genera (16%)include: Cryptomeria, Metasequoia, Pseudotsuga,Pterocarya, Carya, Raphidephis, Sapinium, Punica,Campsis and Liquidambar.

The Holocene period vegetation consists of33 genera, of which Pinus, Abies, Picea, Quercus,Carpinus and Alnus are particularly import-ant. Two extinct genera (6%) are Carya andLespidobalanus.

Changes in the dominant vegetation are recog-nized over the geological periods. Conifers arepredominant (100%) from the Permian to theJurassic, and the dominance of conifers (62%)continued up to the Cretaceous period. On theother hand, the shift of dominant vegeta-tion from the conifers to dicotyledons is afeature of the Cenozoic era. The Oligoceneperiod vegetation was occupied solely by dicoty-ledons (100%), and the predominance of dicoty-ledons (65–81% for the Miocene period; 50–81%

for the Pleistocene period; and 91% for theHolocene period) has been maintained since theOligocene period (Table 2).

The disappearance of all old conifers of thePermian, Triassic and Jurassic periods (100%)implies the occurrence of catastrophic climaticchanges during those periods. Relatively, a highextinction rate of conifers during the Cretaceousperiod (62%) is also indicative of dramatic envir-onmental changes. The sharp decline of theextinction rate of conifers and dicotyledons fromthe Oligocene period (17%), the Miocene period(32–44%), the Pleistocene period (up to 16%) tothe Holocene period (6%) indicates that favourableenvironmental conditions continued during theCenozoic era (Table 2).

Although the oldest plant fossil in Korea(Neuropteris) dates back to the Carboniferousperiod of the Palaeozoic era, the first coniferthat has survived since the Cretaceous periodof the Mesozoic era from both the ROK andthe DPRK is Pinus. This conifer has thrived asone of the dominant tree species since then.

The oldest dicotyledons on the KoreanPeninsula also date back to the Cretaceousperiod, but many of the present-day floristicgenera indeed date to the Oligocene period.The presence of thermophilous genera, suchas Myrica, Ficus and Hedera, in the Oligoceneperiod at up to four degrees farther north oftheir present distributional limits implies thatthe climate of the Oligocene period was warmerthan that of today. The occurrences of otherwarmth-tolerant trees, e.g. Cinnamomum, Cyclo-balanopsis and Ilex at up to six degrees northof the present range during the Middle Miocene,suggests the prevalence of a climatic ‘optimum’.The continued occurrence of numerous present-day genera since the Oligocene indicates eithera long-term stability of Korean vegetation, alongwith fluctuations within it, or habitat diversitieswhich have ensured plant survival in refugia.

The admixture of evergreen coniferous plants,e.g. Taxus, Abies and Thuja, and deciduousbroad-leaved plants indicates that a temperateclimate in Korea prevailed for much of theMiddle Pleistocene. However, the appearanceof deciduous broad-leaved plants and cryophil-ous evergreen coniferous trees, such as Taxus,Abies, Picea, Tsuga, Pinus and Thuja within theKorean Peninsula, also implies the existence of




cold episodes during the Upper Pleistocene.The presence of cold episodes during the UpperPleistocene may also have caused a generalexpansion at this time of temperate deciduousbroad-leaved plants and cryophilous evergreenconiferous plants, associated with the south-ward expansion of these floras in Korea in suchphases (Kong, 1998).

The decline of some cryophilous genera, suchas Pinus (Haploxylon), Picea, Abies and Larixfrom c. 10 000 years marks the continuedclimatic amelioration since then, along withminor climatic fluctuations during the Holocene.However, the vegetational differences betweenthe western and eastern coasts of Korea arenoticeable. Overall, the continued appearanceof both conifers as well as dicotyledons, despitethe sporadic hiatus of fossil data, indicates theabsence of past catastrophic environmental changesin Korea.

ACKNOWLEDGMENTS

I would like to thank D. Watts, J.R. Flenley andtwo anonymous reviewers for valuable commentson the manuscript, and H.S. You for preparationof the figure.

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