correlation of pollen sequences in the neogene palynofloristic regions of china

Palaeoworld 15 (2006) 77–99

Research paper

Correlation of pollen sequences in the Neogenepalynofloristic regions of China

Wei-Ming Wang ∗Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China

Received 1 July 2005; received in revised form 15 March 2006; accepted 31 March 2006Available online 26 May 2006

Abstract

As a result of substantial geographic variations and changing environments, as well as the evolutionary development of the plantkingdom itself, palynofloras in the Neogene of China vary over time and space. This paper gives a correlation of pollen sequences inthe formerly established Neogene palynofloristic regions. It is followed by a detailed description of the pollen zones (assemblages)in the different regions.

The following pollen zones are identified: Betulaepollenites-Betulaceoipollenites-Piceaepollenites (Early Miocene),Caryapollenites-Pinaceae-Sporotrapoidites (late Early Miocene–early Middle Miocene), Caryapollenites-Pinaceae-Ephedripites(late Middle Miocene), Ulmipollenites-herbaceous pollen-Sporotrapoidites (Late Miocene–earliest Pliocene), and Persicariopollis-Chenopodipollis-Magnastriatites (Pliocene) zones in the East China region; Tricolporopollenites-Zonocostatites-Chenopodipollis-Graminidites (early Early Miocene), Polypodiisporites-Florschuetzia-Fonertrites spinosus (syn. Echitricolporites)-Casuarinidites(late Early Miocene–early Middle Miocene), Cupuliferoipollenites-Ilexpollenites-Dicolpopollis-Liquidambarpollenites (lateMCCAMM(P©

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iddle Miocene–Late Miocene), and Cupuliferoipollenites-Polypodiaceaesporites-Chenopodipollis (Pliocene) zones in theoastal Southeast China region; Piceaepollenites-Ulmipollenites-Chenopodipollis (Early Miocene), Potamogetonacidites-henopodipollis-Pinaceae (Middle Miocene), Chenopodipollis-Quercoidites-Ulmipollenites (Late Miocene), and Chenopodipollis-rtemisiaepollenites (Pliocene) zones in the Inland China region; Quercoidites-Betulaceae-Pinuspollenites (late Earlyiocene–early Middle Miocene), Quercoidites-Pinus-Rhododendron (late Middle Miocene), and Quercus-Salix (Lateiocene–earliest Pliocene) in the Southwest China region; Abies-Picea-Pteris (Early Pliocene) and Pinaceae-herbaceous pollen

Late Pliocene) zones in the Qinghai-Xizang region; monolete spores-Quercoidites-herbaceous pollen zone (Early Pliocene) andinaceae-Quercus pollen (Late Pliocene) zones in the Hengduan Mts region.2006 Published by Elsevier Ltd on behalf of Nanjing Institute of Geology and Palaeontology, CAS.

eywords: Pollen zones; Palynofloristic regions; Correlation; Neogene; China

. Introduction

Palynological data accumulated over the last threeecades have greatly broadened knowledge on thehanges of Neogene floras in China. Due to its largeerritory and unevenly developed environmental back-

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ground, detailed correlation among the Neogene pollensequences in the different regions of China was once dif-ficult. Recent progress in regional chronostratigraphicand other related studies have made a systematic studypossible.

The upsurge of Neogene palynology in China reflectsthe development of the oil industry. There were onlylimited reports before the 1970s (Song, 1958, 1959;Ling, 1965). With ever-expanding oil exploration, along

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78 W.-M. Wang / Palaeoworld 15 (2006) 77–99

with advances in related disciplines, Neogene palynol-ogy, as an effective tool in stratigraphic correlation anddivision, quickly became established in some large petro-liferous basins (Sun et al., 1981; Song et al., 1985;Zhu et al., 1985; Guan et al., 1989). The Neogene ofChina is mainly terrestrial with isolated outcrops. Conse-quently, many studies have concentrated on the detailedevaluation of available outcrops and core profiles fortransregional correlations, using palaeomagnetic andother related methods in the oil-gas-bearing areas andsome representative profiles (CGTRP, 1992; Wang andDeng, 2005; Wu et al., 2006). Under the organiza-tion of the National Stratigraphy Committee, combinedefforts have been made to establish the Neogene strati-graphic chart of China, mainly based on fossil mam-mal and palaeomagnetic evidence (National StratigraphyCommittee, 2002; Wang and Deng, 2005). Addition-ally, there are also some syntheses, such as those on theMiocene floristic regions of China (Song et al., 1981)and East Asia (Song et al., 1984), the Neogene paly-nological sequence of Northern China (Liu, 1988), andpalaeofloristic regions and palaeoclimatic implicationsof Neogene palynofloras in China (Wang, 1994; Sun andWang, 2005).

This paper provides a basic correlation of pollensequences in the formerly established Neogene paly-nofloristic regions. General descriptions of pollen zonesand their regional relationships with other pollen data aregiven for each region. Meanwhile, some representativepalynomorphs in the pollen floras are also illustrated.

boundary. Pollen floras in the Late Paleogene to EarlyNeogene period display a gradual change in composi-tion. Most of the types vary quantitatively, while firstand last appearances occur over a period of time ratherthan exactly on the boundary (Song et al., 1985). How-ever, there are still some regional features around theboundary that are more or less recognizable and becomemore and more distinct with time. Changes around thePaleogene/Neogene boundary are mostly related to theevolution and diversification of the plants (Leopold,1969; Song et al., 1985; Wang, 1999) and associatedclimatic changes (Zhao et al., 1982; Sun et al., 1984;Wang and Zhang, 1990). There is evidence to suggestthat major climate events usually occur worldwide, inspite of possible local modifications. The Oligoceneclimate experienced gradual cooling, which partlycontinued into the Early Miocene (Shackleton, 1984).This transition is well-documented in the pollen florasof Northern China (Liu, 1988; Wang and Zhang, 1990).

Because of the restricted nature of terrestrial deposits,stratigraphic correlation between and within the differentregions has been extremely important. The Neogene ofJapan has proved critical because it is largely composedof marine or coastal deposits with good outcrops andestablished floral data. Therefore, the floral sequenceproposed by Tanai (1961) and related pollen zones (Sato,1960, 1963; Yamanoi, 1989) were once widely cited(e.g. Zheng, 1982; Gan, 1982; Hu and Wang, 1985).According to Tanai’s early results (Tanai, 1961), theNeogene floras of Japan include earliest Early Miocene

2. Basic features

The Neogene pollen floras were, in many respects,transitional between those of the Paleogene and those oftoday. This was largely a result of major environmentalvariations during that period, and some accompanyingevolutionary processes in the plant kingdom itself.Physical factors causing the changes mainly includeincreased worldwide climatic fluctuations (Frakes,1979; Berger, 1982; Shackleton, 1984; Kennett and vonder Borch, 1985), the development of modern topogra-phy and the formation of the East Asian Monsoon (Wu,1980; Wang, 1991; An et al., 2001; Sun and Wang,2005). In the Neogene pollen floras, certain originalPaleogene types, including those widely distributedtropical and subtropical elements, gradually becameextinct, while others became largely restricted to lowerlatitudes. Conversely, a number of newly evolved types,such as the dicotyledonous herbs, flourished.

Basically, there is no key element in the pollen florasthat can be used to determine the Paleogene/Neogene

Ainoura-type, Early Miocene Aniai-type, MiddleMiocene Daijima-type, Late Miocene–Early PlioceneShinjo-type and Late Pliocene Akashi-type floras. Thedetailed age assignments of these floral types were lateradjusted in line with advances in chronology (Tanai andUemura, 1991), that is the age of the Ainoura-type wasdetermined to be Oligocene and that of the Daijima-typeto be late Early Miocene–earliest Middle Miocene.Other efforts include direct comparison between Neo-gene pollen floras of Northern China and Japan (Wangand Yamanoi, 1996; Wang and Zhou, 1999; Wang et al.,2001).

The lower boundary of the Quaternary is another sub-ject of palynological study. Climatic and floral changesare generally taken as a key criterion for the division ofthe Pliocene and Pleistocene. As the late Cenozoic cli-mate deteriorated in a stepwise fashion, there are stillcontroversies over the detailed placement of the Plio-Pleistocene boundary (Ogg, 2004; Wang et al., 2004).The Matuyama/Gauss boundary is stratigraphically dis-tinguishable in China, and also represents a major tran-sitional stage for the floras (Liu et al., 2002). It is usually

W.-M. Wang / Palaeoworld 15 (2006) 77–99 79

Fig. 1. Correlation of Neogene pollen sequences in the different palynofloristic regions of China (stratigraphic chart after National StratigraphyCommittee, 2002; Gradstein et al., 2004).

adopted as the Plio-Pleistocene boundary by geologistsand also used in this paper.

Nomenclature for pre-Quaternary fossil pollen andspores in China is mainly based on organ and formgenera established by Potonie (1956, 1960, 1970). Insome cases, the natural classification system is adopted,mainly in the pollen floras since the middle Miocene.Appendix A provides an alphabetic list of all morpho-taxa in the text with their affinity.

Locality data on Neogene floras of China mostlyrefer to some earlier studies (Wang, 1994, p. 240,Fig. 1, p. 241, Table 1; Sun and Wang, 2005, p. 183,Fig. 1, pp. 188–191, Table 1). The division of the Neo-gene palynofloristic regions follows the author’s formerwork (Wang, 1994). At the beginning of the Neogene,the latitudinal zonation of the palynofloristic regionsdeveloped during the Paleogene was replaced by fournew paleofloristic regions, that is East China, CoastalSoutheast China, Inland China and Southwest Chinapaleofloristic regions (Wang, 1994, p. 244, Fig. 2).

Major variations in this floral pattern commenced in thelate Middle Miocene. The Inland China palynofloris-tic region expanded eastwards into the area of InnerMongolia, and the Qinghai-Xizang and Hengduan Mtspalynological regions formed in the existing South-west China palynofloristic region (Wang, 1994, p. 245,Fig. 3).

3. Correlation of pollen sequences

A correlation of pollen sequences in the Neo-gene palynofloristic regions is made in Fig. 1, underthe framework of the international Neogene strati-graphic chart (Gradstein et al., 2004) and the Chi-nese regional Neogene stratigraphic chart (NationalStratigraphy Committee, 2002). Neogene pollen florasin China are generally rich in types (Figs. 2–11), andvary from place to place. Though the pollen zones men-tioned in the paper are largely based on some selecteddata, their time ranges are still regionally changeable.


Fig. 2. (All figures×800 unless otherwise stated): (A) Undulatisporites microverucosus Li, 1983 Yalong, Guangxi; Miocene; (B and G) Multin-odisporites taizhouensis Liu, 1981 Yalong, Guangxi; Miocene; (C) Obtusisporites sp. Bohai Sea; Quaternary Pingyuan Formation (×1000); (Dand H) Pterisisporites undulatus Sung et Zheng, 1976 Yalong, Guangxi; Miocene; (E) Pterisisporites fatangularis Liu, 1981 Yalong, Guangxi;Miocene; (F) Sphagnumsporites stereoides (Potonie et Ven.) Raatz, 1937 Bohai Sea; Lower Miocene Guantao Formation (×1000); (I) Pterisis-porites zonatus Song et Lee, 1976 Yalong, Guangxi; Miocene; (J) Undulatisporites undulapolus Brenner, 1963 Yalong, Guangxi; Miocene; (Kand L) Undulatisporites divergens Wang (1989) Yalong, Guangxi; Miocene; (M) Polypodiaceaesporites haardti (Potonie et Ven.) Thiergart, 1938Zhaotong, Yunan; Pliocene; (N and W) Polypodiisporites communicus Song et Li, 1985 (N) Bohai Sea; Middle Miocene-Pliocene MinghuazhenFormation (×500); (W) East China Sea; Pliocene Santan Formation; (O) Polypodiisporites tenellis (Krutzsch) Song et Zhong, 1984 East China Sea;Pliocene Santan Formation; (P, T and V) Polypodiisporites favus Potonie (1956) (P) Bohai Sea; Middle Miocene-Pliocene Minghuazhen Formation(×500); (T) Kaiyuan, Yunnan; Neogene Xiaolongtan Formation; (V) Bohai Sea; Quaternary Pingyuan Formation (×1000); (Q) Schizaeoisporiteslaevigatiformis (Bolch.) Gao et Zhao, 1976 Yalong, Guangxi; Miocene; (R) Polypodiaceoisporites sp. Yalong, Guangxi; Miocene; (S) Polypodium sp.Zhaotong, Yunnan; Pliocene; (U) Pterisisporites tuberus Song, Li et Zhong, 1986 Yalong, Guangxi; Miocene; (X) Schizaeoisporites dongyingensisKe et Shi, 1978 Yalong, Guangxi; Miocene; (Y) Polypodiisporites sp. Bohai Sea; Lower Miocene Guantao Formation (×1000).


Fig. 3. (All figures ×800 unless otherwise stated): (A) Polypodiaceoisporites sp. Yalong, Guangxi; Miocene; (B) Retitriletes sp. Bohai Sea;Quaternary Pingyuan Formation (×1000); (C) Lycopodiumsporites oligocenicus (Krutzsch) Ke et Shi, 1978 Bohai Sea; Middle Miocene-PlioceneMinghuazhen Formation (×1000); (D and E) Lycopodiumsporites neogenicus (Krutzsch) Ke et Shi, 1978 Bohai Sea; (D) Lower Miocene. GuantaoFormation (×1000); (E) Middle Miocene-Pliocene Minghuazhen Formation (×1000); (F) Deltoidospora regularis (Pflug) Song et Zheng, 1981Yalong, Guangxi; Miocene; (G) Lygodiumsporites meximus (Pflug) Song et Hu, 1985 Bohai Sea; Lower Miocene Guantao Formation (×500); (H)Osmundacidites wellmanii Couper, 1953 Bohai Sea; Quaternary Pingyuan Formation; (I) Toroisporis (Divitoroisporis) cf. zeitzensis Krutzsch, 1959Yalong, Guangxi; Miocene; (J) Deltoidospora adriensis (Pot. et Gell.) Song et Zheng, 1981 Yalong, Guangxi; Miocene; (K) Namlingspora hubeiensis(Ma) Song et Zhong, 1984 East China Sea; Miocene Hailongjing Formation; (L, O and P) Magnastriatites hawardi Germeraad, Hopping et Muller,1968 Bohai Sea; (L and P) Lower. Miocene Guantao Formation (×500); (O) Middle Miocene-Pliocene Minghuazhen Formation (×500). (M)Crassoretitriletes vanraadshooveni Germeraad, Hopping et Muller, 1968 East China Sea; Pliocene Santan Formation (×500); (N) Magnastriatitescf. granulastriatus Liu, 1985 Bohai Sea; Lower Miocene Guantao Formation (×500).


Fig. 4. (All figures ×500 unless otherwise stated): (A and K) Podocarpidites minutus Wang (1978); (A) Zhangpu, Fujian; Neogene Fotan Group;(K) Yalong, Guangxi; Miocene; (B) Dacrydiumites fotanensis Zheng (1987) Zhangpu, Fujian; Neogene Fotan Group; (C) Pinuspollenites labdacusf. minor (Pot.) Potonie, 1958 Easter China Sea; Miocene Hailongjing Formation; (D) Pinuspollenites taedaeformis (Rakl.) Ke et Shi, 1978 BohaiSea; Lower Miocene Guantao Formation; (E and F) Pinuspollenites labdacus f. maxinus (Pot.) Potonie, 1958 Bohai Sea; Lower Miocene GuantaoFormation; (G) Podocarpidites elongatus Song et Zheng, 1981 Bohai Sea; Lower Miocene Guantao Formation; (H) Tsugaepollenites igniculusf. minor Potonie, 1951 Bohai Sea; Lower Miocene Guantao Formation; (I) Abietineaepollenites microalatus f. minor Potonie, 1951 Bohai Sea;Quaternary Pingyuan Formation; (J) Dacrydiumites elegans Zhang, 1981 Leqiong Area, Guangdong; Neogene. (L) Podocarpidites nageiaformis(Zakl.) Krutzsch, 1971 Bohai Sea; Lower Miocene Guantao Formation; (M) Piceaepollenites praemarianus (Krutzsch) Song (1988) Zhaotong,Yunnan; Pleistocene; (N and O) Ephedripites (Distachyapites) fusiformis (Shakhmundes) Krutzsch, 1970 Yalong, Guangxi; Miocene (×800); (P)Tsugaepollenites igniculus f. major Potonie, 1958 East China Sea; Miocene Yuquan Formation; (Q and R) Taxodiaceaepollenites hiatus (Pot.) Kremp,1949; (Q) East China Sea; Miocene Yuquan Formation (×800); (R) Bohai Sea; Lower Miocene Guantao Formation (×1000); (S) Sequoiapollenitespolyformosus Thiergart, 1938 Bohai Sea; Lower Miocene Guantao Formation (×1000); (T) Piceaepollenites tobolicus (Panova) Ke et Shi, 1978Bohai Sea; Lower Miocene Guantao Formation; (U) Piceaepollenites gigantea Wang, 1981 Bohai Sea; Quaternary Pingyuan Formation; (V)Ephedripites megafusiformis Ke et Shi, 1978 Bohai Sea; Lower Miocene Guantao Formation (×1000).


Fig. 5. (All figures ×500 unless otherwise stated): (A and H) Piceaepollenites tobolicus (Ponova) Ke et Shi, 1978 Zhaotong, Yunnan; (A) Pliocene,H. Pleistocene; (B) Tsugaepollenites spinulosus (Krut.) Ke et Shi, 1978 Bohai Sea; Lower Miocene Guantao Formation; (C) Ephedripites (Dis-tachyapites) fusiformis (Shakhmundes) Krutzsch, 1970 Yalong, Guangxi; Miocene (×800); (D) Abiespollenites sp. 1 (cf. Abies nephrolepis)Zhaotong, Yunnan; Pleistocene; (E) Abiespollenites sp. 2 (cf. Abies sutchuensis) Zhaotong, Yunnan; Pleistocene; (F) Ephedripites (Ephedrip-ites) notensis (Cookson) Krutzsch, 1961 Yalong, Guangxi; Miocene (×800); (G and I) Abiespollenites sibirieiformis (Zakl.) Krutzsch, 1971 BohaiSea; (G) Quaternary Pingyuan Formation; (I) Lower Miocene Guantao Formation; (J) Piceaepollenites sp. Zhaotong, Yunnan; Pleistocene.


Fig. 6. (All figures × 800 unless otherwise stated): (A) Platycaryapollenites sp. Yalong, Guangxi; Miocene; (B) Platycaryapollenites cf. dongyin-gensis Ke et Shi, 1978 Yalong, Guangxi; Miocene; (C) Engelhardtia sp. 1 Zhaotong, Yunnan; Pliocene; (D) Engelhardtia sp. 2 Zhaotong, Yunnan;Pliocene; (E) Ulmoideipites krempii Anderson, 1960 Yalong, Guangxi; Miocene; (F and G) Ulmoideipites tricostatus Anderson, 1960 Yalong,Guangxi; Miocene; (H and I) Myrica esculenta Buch.-Ham. Zhaotong, Yunnan, Pliocene; (J) Betulaepollenites claripites (Wodeh.) Sung et Tsao,1976 Bohai Sea, Lower Miocene Guantao Formation (×1000); (K) Betulaceoipollenites bituitus (Pot.) Potonie, 1951 Bohai Sea; Lower MioceneGuantao Formation (×1000); (L and W) Alnipollenites verus (Pot.) Potonie, 1934; (L) East China Sea; Miocene Hailongjing Formation; (W) BohaiSea; Lower Miocene Guantao Formation (×1000); (M) Symplocoipollenites vestibulum (Pot.) Potonie (1960) Bohai Sea; Quaternary PingyuanFormation; (N) Corylus ferox Wall. Zhaotong, Yunnan; Pleistocene; (O) Myriophyllum verticillatum L. Zhaotong, Yunnan; Pleistocene; (P and U)Betulaepollenites plicoides (Zakl.) Sung et Tsao, 1976 Bohai Sea; (P) Lower Miocene Guantao Formation (×1000); (U) Quaternary PingyuanFormation (×1000); (Q and R) Micrechites sp. Zhaotong, Yunnan; Pliocene; (S) Carpinipites spackmonii (Trav.) Zhou, 1981 Bohai Sea; LowerMiocene Guantao Formation (×1000); (T) Myriophyllumpollenites minutus Nagy, 1985 Bohai Sea; Quaternary Pingyuan Formation (×1000); (V)Rhoipteapollis chilianthoides Zheng, 1985 East China Sea; Pliocene Santan Formation; (X) Alnipollenites metaplasmus (Pot.) Potonie (1960) BohaiSea; Lower Miocene Guantao Formation (×1000); (Y) Zelkova sp. Zhaotong, Yunnan; Pliocene; (Z) Zelkovaepollenites cf. potonie Nagy, 1969East China Sea; Miocene Hailongjing Formation; (AA) Ulmoideipites sp. (Planera aquatica J.F. Gmel.) Zhaotong, Yunnan; Pliocene; (BB and GG)Ulmoideipites neogenicus Guan, 1989 Bohai Sea; Quaternary Pingyuan Formation (×1000); (CC) Betulaceoipollenites cf. bituitus (Pot.) Potonie,1951 Bohai Sea; Lower Miocene Guantao Formation (×1000); (DD and EE) Carya tonkinensis Lee Zhaotong, Yunnan; Pleistocene; (FF) Juglans-pollenites rotundus Ke et Shi, 1978 Bohai Sea; Lower Miocene Guantao Formation; (HH) Pterocaryapollenites stellatus (Pot.) Raatz, 1938 BohaiSea; Lower Miocene Guantao Formation (×1000); (II) Juglanspollenites verus Raatz, 1939 Bohai Sea; Middle Miocene-Pliocene MinghuazhenFormation; (JJ) Multiporopollenites rariporus Guan, 1989 Bohai Sea; Quaternary Pingyuan Formation (×1000).


Fig. 7. (All figures ×800 unless otherwise stated): (A) Castanea sp. Zhaotong, Yunnan; Pleistocene; (B) Cupiliferoipollenites oviformis Potonie(1960) Kaiyuan, Yunnan; Neogene Xiaolongtan Formation; (C and D) Cupuliferoipollenites pusillus (Pot.) Potonie (1960); (C) Yalong, Guangxi;Miocene; (D) Kaiyuan, Yunnan; Neogene Xiaolongtan Formation; (E) Quercus sp. Zhaotong, Yunnan; Pleistocene; (F) Philadelphus sp. Zhaotong,Yunnan; Pliocene; (G) Cyrillaceaepollenites miniporus Zheng, 1985 East China Sea; Miocene Yuquan Formation; (H) Tricolpopollenites oblatusWang, 1985 East China Sea; Miocene Hailongjing Formation; (I) Quercoidites microhenrici (Pot.) Potonie, 1950 Kaiyun, Yunnan; XiaolongtanFormation; (J) Quercoidites henrici (Pot.) Potonie, Thomson et Thiergart, 1950 Bohai Sea; Lower Miocene Guantao Formation (×1000); (Kand L) Trilobapollis ellipticus Sun, Kong et Li, 1980; (K) East China Sea; Miocene Hailongjing Formation; (L) Yalong, Guangxi; Miocene.(M) Rutaceoipollis minireticulatus Wang (1989) Yalong, Guangxi; Miocene; (N) Florschuetzia trilobata Germeraad, Hopping et Muller, 1968Zhujiangkon, Guangdong; Miocene Hanjiang Formation; (O and P) Florschuetzia levipoli Germeraad, Hopping et Muller, 1968 Beibuwan, Guangxi;(O) Miocene Jiaowei Formation; (P) Pliocene Wangloujiao Formation; (Q and W) Distyliumpollis trilobatus Wang (1989) Yalong, Guangxi; Miocene;(R and S) Rutaceoipollis ovatus Sung et Tsao, 1980 Bohai Sea; Lower Miocene Guantao Formation (×1000); (T) Rutaceoipollis sp. Yalong, Guangxi;Miocene; (U and V) Hydrocotaepites pachydermus (Sun et al.) Zheng, 1985 East China Sea; (U) Pliocene Santan Formation; (V) Miocene YuquanFormation; (X) Nyssapollenites microreticulatus Zheng, 1989 Bohai Sea; Lower Miocene Guantao Formation; (Y) Morinda umbellata L. Zhaotong,Yunnan; Pleistocene; (Z and EE) Sporotrapoidites medius Guan, 1985 East China Sea; Miocene Hailongjing Formation; (AA, BB and FF–HH)Sporotrapoidites erdtmanii (Nagy) Nagy, 1985 Bohai Sea; (AA, BB and GG) Lower Miocene Guantao Formation; (BB and GG) (×1000); (FF andHH) Middle Miocene-Pliocene Minghuazhen Formation; 34 (×1000); (CC and DD) Sporotrapoidites minor Guan, 1985 East China Sea; MioceneHailongjing Formation; (II) Sporotrapoidites weiheensis (Sun et Fan) Guan, 1985 Bohai Sea; Middle Miocene-Pliocene Minghuazhen Formation.


Fig. 8. (All figures ×800 unless otherwise stated): (A) Proteacidites spiniformis Ke et Shi, 1978 East China Sea; Miocene Hailongjing Formation;(B) Symplocoipollenites latiporis (Thomson et Pflug) Zheng, 1985 East China Sea; Miocene Yuquan Formation; (C) Nyssapollenites microreticulatusZheng, 1989 Bohai Sea; Lower Miocene Guantao Formation (×1000); (D) Corsinipollenites triangulus (Zakl.) Ke et Shi, 1978 Bohai Sea; QuaternaryPingyuan Formation; (E, M and N) Lonicerapollis gallwitzii Krutzsch, 1962 5. East China Sea; Miocene Hailongjing Formation; (M and N) BohaiSea; Lower Miocene Guantao Formation (×1000); (F) Lonicerapollis cf. triletus Zheng, 1985 Bohai Sea; Middle Miocene-Pliocene MinhuazhenFormation; (G) Rhoipteapollis chilianthoides Zheng, 1985 East China Sea; Pliocene Santan Formation (×1000); (H) Retitricolporites conspicuusLiu, 1981 Yalong, Guangxi; Miocene; (I) Tiliaepollenites pseudoinstructus (Mai) Ke et Shi, 1978 Bohai Sea; Lower Miocene Guantao Formation(×1000); (J) Tiliaepollenites indubitabilis Potonie, 1953 East China Sea; Pliocene Santan Formation; (K) Tiliaepollenites instructus Potonie, 1951Bohai Sea; Lower Miocene Guantao Formation; (L) Lonicera japonica Thunb. Zhaotong, Yunnan; Pliocene; (O) Ranunculacidites vulgaris Song etG.X. Li Bohai Sea; Middle Miocene-Pliocene Minghuazhen Formation; (P) Corsinipollenites subrotundus Zheng, 1989 Bohai Sea; Lower MioceneGuantao Formation (×1000); (Q) Corsinipollenites triangulus (Zakl.) Ke et Shi, 1978 Bohai Sea; Lower Miocene Guantao Formation (×1000); (R)Lonicerapollis rarus Zheng, 1985 East China Sea; Miocene Hailongjing Formation.


Fig. 9. (All figures ×800 unless otherwise stated): (A and B) Operculumpollis operculatus Sun, Kong et Li, 1980 East China Sea; Miocene Hai-longjing Formation; (C) Echitricolporites sp. Bohai Sea; Quaternary Pingyuan Formation (×500); (D) Margocolporites minor Zheng, 1985 EastChina Sea; Pliocene Santan Formation; (E) Ilexpollenites membranous Sun, Kung et Li, 1980 Kaiyuan, Yunnan; Neogene Xiaolongtan Formation;(F) Retimultiporopollenites minor Guan, 1985 East China Sea; Pliocene Santan Formation; (G) Cyrillaceaepollenites megaexactus (Pot.) Potonie(1960) Bohai Sea; Lower Miocene Guantao Formation (×1000); (H) Rutaceoipollis ovatus Sung et Tsao, 1980 Bohai Sea; Lower Miocene Guan-tao Formation (×1000); (I) Callicarpapollis firmexinus Wang (1989) Yalong, Guangxi; Miocene; (J) Abutilonacidites bohaiensis Guan et Zheng,1989 Bohai Sea; Middle Miocene-Pliocene Minghuazhen Formation (×500); (K) Rutaceoipollis parviporus Zheng, 1985 East China Sea; MioceneHailongjing Formation; (L) Aceripollenites communis Zheng, 1989 Bohai Sea; Lower Miocene Guantao Formation (×1000); (M) Retitricolpitesmatauraensis (Couper) Song et Zheng, 1981 East China Sea; Miocene Hailongjing Formation; (N) Labitricolpites cf. microgranulatus Ke et Shi,1978 East China Sea; Pliocene Santan Formation; (O) Faguspollenites sp. East China Sea; Miocene Yunquan Formation; (P) Nyssapollenitesmicroreticulatus Zheng, 1989 Bohai Sea; Lower Miocene Guantao Formation (×1000); (Q) Cruciferaeipites minor Zheng, 1989 Bohai Sea; Quater-nary Pingyuan Formation (×1000); (R) Symplocospollenites communis Ke et Shi, 1978 Yalong, Guangxi; Miocene; (S) Micrechites sp. Zhaotong,Yunnan; Pliocene; (T) Micrechites lachnocarpa Tsiang Zhaotong, Yunnan; Pliocene; (U) Myrtaceidites reticulatus Wang (1989) Yalong, Guangxi;Miocene; (V) Gothanipollis longjingensis Zheng, 1985 East China Sea; Miocene Hailongjing Formation; (W) Loranthacites rhonmbiformis Wang(1989) Yalong, Guangxi; Miocene; (X) Retitricolpites sp. East China Sea; Miocene Yuquan Formation; (Y) Liquidambarpollenites orientaliformisNagy, 1969 Bohai Sea; Lower Miocene Guantao Formation; (Z) Liquidambarpollenites pachydermus Ke et Shi, 1978 Bohai Sea; Lower MioceneGuantao Formation (×1000); (AA) Liquidambarpollenites stigmosus (Pot.) Raatz, 1938 Bohai Sea; Middle Miocene-Pliocene Minghuazhen Forma-tion (×1000); (BB) Celtispollenites dongyingensis Ke et Shi, 1978 Bohai Sea; Lower Miocene Guantao Formation (×1000); (CC) Juglanspollenitesverus Raatz, 1939 Bohai Sea; Middle Miocene-Pliocene Minghuazhen Formation (×1000); (DD and EE) Caryapollenites simplex (Pot.) Raatz,1937; (DD) Kaiyuan, Yunnan; Xiaolongtan Formation; (EE) Bohai Sea; Lower Miocene Guantao Formation (×1000).


Fig. 10. (All figures ×800 unless otherwise stated): (A) Lonicerapollis granulatus Ke et Shi, 1978 Yalong, Guangxi; Miocene; (B) Tricolpopollenitessp. East China Sea; Pliocene Santan Formation; (C) Rhoipites pseudodolium Song et Zheng, 1981 Yalong, Guangxi; Miocene; (D) Labitricolpitesstenosus Ke et Shi, 1978 Bohai Sea; Lower Miocene Guantao Formation (×1000); (E) Yalongipollis gracilis Wang (1989) Yalong, Guangxi; Miocene;(F) Fupingopollenites wackersderfensis (Thiele-Pfeiffer) Liu, 1985 Yalong, Guangxi; Miocene; (G) Fupingopollenites minutus Liu, 1985 Yalong,Guangxi; Miocene; (H) Aceripollenites striatus (Pflug) Thiele-Pfeiffer, 1980 East China Sea, Miocene Yuquan Formation; (I) Faguspollenitessp. East China Sea; Miocene Yuquan Formation; (J) Retitrescolpites pellucidus Zheng, 1989 Bohai Sea; Middle Miocene-Pliocene MinghuazhenFormation (×500); (K) Marginipollis robustus (Playford) Zheng, 1984 Zhangpu, Fujiang; Neogene Fotan Group; (L) Marginipollis elegans Zheng,1984 Zhangpu, Fujiang; Neogene Fotan Group; (M and N) Yalongipollis medius Wang (1989) Yalong, Guangxi; Miocene; (O) Rutaceoipollis cf.reticulatus Song et Li, 1982 Yalong, Guangxi; Miocene; (P and Q) Scabiosapollis radiostratus Song, 1985 (P) Bohai Sea; Middle Miocene-PlioceneMinghuazhen Formation (×500); (Q) East China Sea, Pliocene Santan Formation (×500); (R and S) Abutilonacidites bohaiensis Guan et Zheng,1989 Bohai Sea; Middle Miocene-Pliocene Minghuazhen Formation; (T) Echitricolporites sp. Yalong, Guangxi; Miocene.


Fig. 11. (All figures ×800 unless otherwise stated); (A–D) Reevesiapollis triangulus (Mamczar) Krutzsch, 1980 (A and B) East China Sea; PlioceneSantan Formation; C and D) Yalong, Guangxi; Miocene; (E, F and U) Artemisiaepollenites sellularis Nagy, 1969; (E) Yalong, Guangxi; Miocene;(F and U) Bohai Sea; Quaternary Pingyuan Formation (×1000); (G) Artemisiaepollenites leatus Zheng, 1989 Bohai Sea; Lower Miocene GuantaoFormation; (H) Reevesiapollis siameniformis Guan et Zheng, 1989 Bohai Sea; Lower Miocene Guantao Formation; (I) Cichorieacidites ixeriformisZheng, 1989 Bohai Sea; Middle Miocene-Pliocene Minghuazhen Formation; (J and K) Chenopodipollis minor Song, 1985 Bohai Sea; (J) LowerMiocene Guantao Formation; (K) Quaternary Pingyuan Formation; (L) Dicolpopollis kochelii Pflanzi, 1956 Beibuwan, Guangdong; MioceneJiaowei Formation; (M) Euphorbiacidites sp. East China Sea; Quaternary Donghai Group; (N) Persicarioipollis lusaticus Krutzsch, 1962 BohaiSea; Middle Miocene-Pliocene Minghuazhen Formation (×1000); (O) Percicarioipollis communis Ke et Shi, 1978 Bohai Sea; Quaternary PingyuanFormation; (P) Caryophyllidites tenius Song et Zhu, 1985 Bohai Sea; Quaternary Pingyuan Formation (×1000); (Q) Amaranthaceae Zhaotong,Yunnan; Pleistocene; (R) Umbelliferaepites microreticulatus Zheng, 1985 East China Sea; Quaternary Donghai Group; (S) Umbelliferaepites cf.ovatus (Vank. et Kar.) Zheng, 1985 East China Sea; Pliocene Santan Formation; (T) Umbelliferaepites bupleurumiformis Zheng, 1985 East ChinaSea; Pliocene Santan Formation; (V and W) Nymphaeacidites minor (Nagy) Zheng, 1985 Bohai Sea; (V) Middle Miocene-Pliocene MinghuazhenFormation (×1000); (W) Quaternary Pingyun Formation (×1000); (X) Pieris ovalifolia D. Don. Zhaotong, Yunnan; Pleistocene; (Y) Ericipitesericius Potonie (1960) East China Sea; Pliocene Santan Formation; (Z) Margocolporites cristoides Zheng, 1985 East China Sea; Pliocene SantanFormation; (AA and BB) Graminidites subtiliglobosus (Trevisan) Krutzsch, 1970 Bohai Sea; Quaternary Pingyun Formation (×1000); (CC)Graminidites pseudograminens Krutzsch, 1970 Bohai Sea; Lower Miocene Guantao Formation (×1000); (DD) Beaupreaidites sp. Bohai Sea;Middle Miocene-Pliocene Minghuazhen Formation (×1000); (EE) Liliacidites sp. 1 Bohai Sea; Lower Miocene Guantao Formation (×1000); (FF)Liliacidites sp. 2 East China Sea; Pliocene Santan Formation.


Based on the pollen sequences in different regions,it seems that the pollen floras were more or less evenlydeveloped in the early Neogene. Intensified diversifica-tion started from the Middle Miocene, mostly caused byenvironmental changes. The development of Neogeneenvironments in China was influenced by various factors.With the worldwide climatic deterioration as major con-trol, some local events also exerted a distinct influence,such as the development of the Himalayan Mountain sys-tem and the formation of the East Asian monsoon (Li etal., 1979; An et al., 2001; Sun and Wang, 2005).

At the beginning of the Neogene, some areas in thewestern part of the Inland China region might still havebeen partly affected by the Tethys climate. For example,the pollen flora from the Early Miocene middle andupper parts of the Jidike Formation in the Kuche Basinis marked by a higher representation of amentiferousplants in types and values, together with a small numberof tropical and subtropical elements such as Palmaeand Trochodendraceae, in addition to a high diversityof gymnospermous taxa (Zhao et al., 1982; Wang etal., 1990). This region was generally dominated byxerophytic plants. The arid climate expanded eastward,and the Inner Mongolia subregion was gradually formedfrom the late Middle Miocene. The pollen flora from thelate Middle Miocene Tongguer Formation, Inner Mon-golia, for example, is represented by about half of theherbaceous pollen in the total dominant angiospermouspollen, including Chenopodipollis, Artemisiaepollen-ites, Graminidites, and Compositoipollenites (syn.

region are mainly based on data from the North Chinapalynofloristic subregion, but comparisons with thepollen floras from the other two subregions are providedas well.

4.1.1. Betulaepollenites-Betulaceoipollenites-Piceaepollenites zone (Early Miocene)

This zone contains a dominant proportion ofangiospermous pollen, a moderate amount of gym-nospermous pollen, and a small number of pterido-phyte spores. In the angiospermous pollen, there ishigh representation for Betulaepollenites (2.8–13.0%)and Betulaceoipollenites (2.8–11.5%), together withsome frequently occurring elements, including Ulmipol-lenites, Zelkovaepollenites, Juglanspollenites, Quer-coidites, Ulmoideipites, Momipites and Alnipollenitesamong the woody plants, and Sporotrapoidites amongthe herbs. The gymnospermous pollen is marked byhigh values for Piceaepollenites (9.8–19.3%), alongwith some representation of Tsugaepollenites, Podocar-pidites, Abiespollenites, Abietineaepollenites, Pinuspol-lenites; and sporadic Taxodiaceaepollenites. Polypodi-aceaesporites and Osmandacidites are well-representedin the pteridophyte spore component. This zone descrip-tion is from the Shangdou-Huade Basin, Inner Mongolia(Wang and Zhang, 1990).

Other comparable pollen floras of the same subre-gion are found in the basaltic inter-bedded deposit atWuluogong, North Hebei Province (Gan, 1982), the lig-nite deposit at Tianzhen, Shanxi Province (Wang, 1978),

Echitricolporites) (Wang, 1990).The pollen floras in the Southwest China paly-

nofloristic region show the major influence of a sec-ond Himalayan movement. As the result, the Qinghai-Xizang and Hengduan Mts palynological regions gradu-ally developed from the existing Southwest China paly-nofloristic region. The alpine pollen floras formed inSouthwest China are consistent with the uplift of the landsince the Middle Miocene. Changes in the East Chinaand Coastal Southeast China palynofloristic regions arenot as distinct as in the other two regions mainly due toits geographic location. However, some general changesare still recognizable, such as increased floral variationand a rise of herbs.

4. Description of pollen zones

4.1. East China palynofloristic region

This region covers the whole of Eastern China andincludes the Northeast China, North China and SouthChina palynofloristic subregions. Pollen zones in the

Drill Hole T103 in Tianchang, Anhui Province (Zhengand Zhang, 1986), and the lower part of the GuantaoFormation, Bohai Sea (Guan et al., 1989).

In the Northeast China subregion, the same periodis represented by the Pinus-Picea-Tsuga-Podocarpusassemblage from the Three-River Plain. It is domi-nated by gymnosperm pollen (77.8%), including Pinus(28.8%), Picea (20.1%), Tsuga (13.7%), Podocarpus(6.7%) and Taxodium, together with some angiospermpollen, such as Liquidambar, Nyssa, Magnolia, Fagusand pteridophyte spores (Xia et al., 1987).

The Quercoidites-Chenopodipollis assemblage fromthe middle part of the Toupo Group at the Toupo Basin,Jiangxi Province (Sun and He, 1987) is representa-tive of the South China subregion. It shows strongpollen domination by woody plants (ca. 70%), includ-ing Quercoidites, Liquidambarpollenites, Faguspollen-ites, Rhoipites, Myrtaceidites, Magnolipollis and Meli-aceoidites among the angiosperms, and Keteleeriaepol-lenites, Abietineaepollenites, Pinuspollenites, Tsugae-pollenites, Laricoidites, Inaperturopollenites among thegymnosperms. Herbaceous pollen grains are also well


represented (ca. 24%), particularly by Chenopodipol-lis, along with sporadic Ranunculaceae, Cruciferae, Eri-caceae, Labiatae and Rubiaceae. Pteridophyte spores(6%) are mainly represented by Polypodiaceae.

4.1.2. Caryapollenites-Pinaceae-Sporotrapoiditeszone (late Early Miocene–early Middle Miocene)

This zone shows a significant increase in the valuesand types of angiosperm pollen (82%). There is areasonable amount of gymnosperm pollen (17%),and a small amount of pteridophyte spores. Somefrequently occurring elements include Sporotrapoidites,Fupingopollenites, Chenopodiaceae, Caryapollenites,Ulmipollenites, Sparganiaceaepollenites, Momipites,Zelkovaepollenites, Tiliaepollenites, Gramineae andAsteraceae within the angiosperms; Piceaepollenites,Abietineaepollenites, Pinuspollenites, Keteleeriaepol-lenites, Podocarpidites, Tsugaepollenites, Ephedripiteswithin the gymnosperms; and Pterisisporites andDeltoidospora within the pteridophytes. This zone isrepresented by the pollen assemblage from the outcropof the Lengshuigou Formation, and in the lower part ofthe Gaoling Group derived from drilling cores in theWeihe Basin, Shaanxi Province (Sun et al., 1980).

Other comparable assemblages in the subregioninclude those from the Shangdou-Huade Basin,Inner Mongolia (Sporotrapoidites-Ulmipollenites +Zelkovaepollenites and Lonicerapollis-Quercoidites-Polypodiaceaesporites assemblages) (Wang and Zhang,1990), the Shanwang Formation of Linqu, ShandongPueH

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lenites, Tsugaepollenites and Keteleeriaepollenites, areall poorly represented. There are a few Chenopodi-aceae, Polygonaceae, Labiatae, Umbelliferae, Rubi-aceae, Gramineae and Asteraceae, among the herbs.Pteridophyte spores are represented by Polypodiaceaes-porites (Sun and He, 1987). Pollen assemblages from theinterbasaltic beds at Tiantai, Xinchang and Shengxianof Zhejiang Province (Zheng, 1982; Wang et al., 1985),and the lignite deposit of Yalong at Yao County, GuangxiProvince (Wang, 1989), are also comparable to the zone.

4.1.3. Caryapollenites-Pinaceae-Ephedripites zone(late Middle Miocene)

The overall representation of angiospermous pollenhas decreased while pollen of the gymnosperms andsome drought-resistant plants have increased corre-spondingly. Percentages for angiospermous pollen, gym-nospermous pollen and pteridophyte spores average 55,43 and 2%, respectively. The main angiosperm elementsare Ulmipollenites, Caryapollenites and Trochodendron,with Artemisia showing its first presence. There arehigh values for Ephedripites within the gymnosperms,while Pinuspollenites, Piceaepollenites and Keteleeriae-pollenites have notable values. The pollen assemblage ofthis zone was reported from the upper part of the GaolingGroup in the drilling cores of the Weihe Basin, ShaanxiProvince (Sun et al., 1980).

This zone is more distinguishable in the NorthChina palynofloristic subregion than in the othertwo subregions in the region. The Magnastriatites-

rovince (Song, 1959; Liu and Leopold, 1992), thepper part of the Guantao Formation, Bohai Sea (Guant al., 1989), and the Huanghai Formation of the Southuanghai Sea Basin (Li et al., 1984).In the Northeast China subregion, the palynologi-

al assemblage from the Three-River Plain is distin-uished by ameniferous plants. There are Betula, Quer-us, Corylus, Ostrya, Castanea, Ulmus, Juglans, Carya,iquidambar and Fagus, among the woody angiosperms,nd Pinus, Podocarpus, Pseudolarix, Tsuga, Cedrus,eteleeria and Taxodium among the gymnosperms. Botherbaceous pollen and pteridophyte spores display lowalues (Xia et al., 1987).

In the South China subregion, the Quercoidites-iquidambarpollenites assemblage from the upper partf the Toupo Group in Toupo Basin, Jiangxi Province,s characterized by the predominance of woody plantsaverage over 90%), 6% pteridophyte spores and lesshan 5% herbaceous pollen (Sun and He, 1987). Amonghe woody plants, Quercoidites and Liquidambarpol-enites are dominant, and others such as Faguspol-enites, Ulmipollenites, Abietineaepollenites, Pinuspol-

Fupingopollenites-Liquidambarpollenites assemblagefrom the lower part of the Minghuazhen Formation,Bohai Sea (Guan et al., 1989) might be comparable tothis zone. A pollen assemblage from the Yaoshan For-mation of Shanwang, Shandong Province (Liu, 1986)shares similar characteristics with the one revealed fromthe Shanwang Formation of the previous zone, and issuggested to represent the top of the latter (Wang et al.,2002). It might partly extend to this zone.

4.1.4. Ulmipollenites-herbaceous pollen-Sporotrapoidites zone (Late Miocene-earliestPliocene)

This zone is dominated by angiosperm pollen, fol-lowed by gymnosperm pollen and pteridophyte spores.Among the angiosperms, woody plants are still the mainelements, and include Betulaceae, Ulmipollenites andJuglandaceae, while herbs show higher values than in theprevious zone. Gymnosperm pollen has also increasedslightly, mainly represented by Pinaceae, together withsome Taxodiaceae. Very low values of Magnastriatitesoccur within the pteridophyte spores. The zone is estab-


lished from the lower part of the upper MinghuazhenFormation, Bohai Sea (Guan et al., 1989).

Other assemblages in the North China palynofloris-tic subregion include those from depths 390–402 ofthe drilling core at Huanghua, Hebei Province (Li andLiang, 1981) and the IV Formation of the YongledianGroup, Weihe Basin, Shaanxi Province (Sun et al., 1980)amongst others.

The Pinus-Betula-Alnus assemblage from the Three-River Plain of the Northeast China subregion might alsorepresent the same stage. It shows relatively high valuesfor gymnosperm pollen, especially Pinus (52.7%).Angiosperm pollen is low in diversity, mainly includingBetula, Alnus, Juglans, Quercus, Ulmus and Pterocaryaamong the woody plants, and Artemisia and Chenopo-diaceae among the herbaceous pollen (Xia et al., 1987).

4.1.5. Persicariopollis-Chenopodipollis-Magnastriatites zone (Pliocene)

This zone is characterized by a dominance ofangiospermous pollen, secondary pteridophyte spores,and low values for gymnospermous pollen. Amongthe angiosperms, there are more herbs than woodyplants with the herb component dominated by Persi-carioipollis and, occasionally Chenopodipollis locally.Other herbs include Graminidites, Artemisiaepollen-ites, Tubulifloridites, and Echitricolporites, while woodytaxa include Ulmipollenites, Salixipollenites, Alnipol-lenites, Caryapollenites, Nyssapollenites, Magnolipol-lis and Liquidambarpollenites. Magnastriatites are

In the South China subregion, the Persicariopollis-Graminidites assemblage from the Shelf Basin of theEast China Sea can be divided into two subassem-blages. The lower Persicariopollis-Graminidites-Retimultiporopollenites subassemblage is distin-guished by high values of herbaceous pollen, includ-ing Persicariopollis, Graminidites, Asteraceae andChenopodiaceae, together with some woody plantsincluding Liquidambarpollenites, Quercoidites,Ulmipollenites, Juglanspollenites, Betulaceae andTaxodiaceae. The upper Persicariopollis-Graminidites-Polypodiaceaesporites subassemblage is characterizedby higher values of Polypodiaceaesporites and gym-nospermous pollen. Among the angiospermous pollen,Ulmaceae, Juglandaceae, Persicariopollis, Gramini-dites, as well as some subtropical elements, have lowervalues, while values for Betulaceae, Chenopodiaceaehave risen (Song et al., 1985).

4.2. Coastal Southeast China floristicpalynofloristic region

4.2.1. Tricolporopollenites-Zonocostatites-Chenopodipollis-Graminidites zone(early Early Miocene)

Representations of angiospermous pollen, pterido-phyte spores and algae, and gymnospermous pollenin this zone are 69.3–93%, 4–27% and less than10%, respectively. Quercoidites (Q. microhenrici),Cupuliferoipollenites (C. oviformis, C. pusillus) are dis-

distinct among the pteridophyte spores, with othercommon elements including Polypodiaceaesporites,Salviniaspora, Lycopodiumsporites, Polypodiidites andOsmundacidites. The gymnospermous pollen mainlyconsists of Pinaceae, and Taxodiaceae. This zone isrecorded from the upper part of the upper MinghuazhenFormation, Bohai Sea (Guan et al., 1989).

Late in the zone, pollen floras in many places showevident fluctuations. For example, the pollen flora ofthe Yushe Basin, Shanxi Province has consistent rep-resentation of Artemisia, Chenopodiaceae, Ulmus andPinaceae pollen. There are also some minor thermophilicelements such as Juglans/Pterocarya and Carya, whichrepeatedly appear and disappear in the pollen sequence.These indicate a decline in temperature about 3.2 Ma,a warm phase about 3.05 Ma and a cooling event about2.5 Ma (Liu et al., 2002). Other reports of the zone mightinclude, for example, Zone I–II of the Datong Basin,Shanxi Province; Zone I–III of the Weixian Basin, HebeiProvince (Tang and Liu, 1984); the upper YanchengFormation of the Northern Jiangsu Province and SouthYellow Sea Basin (Zheng et al., 1981).

tinctive components with high values in the angiosperms,together with Chenopodipollis, Graminidites and Zono-costatites. There are small amounts of Dacrydiumitesand Tsugaepollenites in the gymnospermous pollen, andmore monolete spores than trilete ones in the pterido-phytes (Sun et al., 1981). This zone is recorded in theXiayang Formation of the Beiduwan depression andLeiqiong areas, and the North Continental Shelf of theSouth China Sea.

A similar pollen flora is found in the Zhujiang For-mation of the Zhujiangkou depression and the MeisahanFormation of the Yingge depression in the same area(Sun et al., 1981).

4.2.2. Polypodiisporites-Florschuetzia-Fonertritesspinosus (syn. Echitricolporites)-Casuariniditeszone (late Early Miocene–early Middle Miocene)

This zone is characterized by a dominance ofangiospermous pollen (52–88%), increasing pterido-phyte spores (5.1–37%) and a small amount ofgymnospermous pollen (0–10%). The angiospermouspollen is well represented by Quercoidites (Q. micro-


henrici, Q. minutus), Cupuliferoipollenites (C. ovi-formis), along with some newly recorded types, forexample, Florschuetzia (F. cf. leviooli), Echitricol-porites and Casuarinidites. Pteridophyte spores are bestrepresented by Polypodiaceoisporites, Polypodiisporitesand Polypodiaceaesporites. The gymnosperm compo-nent is composed mainly of Dacrydiumites, with a fewPodocarpidites and Pinuspollenites (Sun et al., 1981).The zone is recorded in the Jiaowei Formation of the Bei-duwan depression and Leiqiong areas and on the NorthContinental Shelf of the South China Sea.

Pollen assemblages from several other sites in theregion might be synchronous with this zone. Theseinclude the lower part of the Hanjiang Formation in theZhujiangkou depression, the Huangliu Formation of theYinggehai depression on the North Continental Shelf ofthe South China Sea (Sun et al., 1981), the Fotan Groupfrom Zhangpu County, Fujian Province (Zheng, 1987,1988), and the “Middle Coal-bearing Bed” in the ShihtiFormation, Taiwan (Ling, 1965; Canright, 1974).

4.2.3. Cupuliferoipollenites-Ilexpollenites-Dicolpopollis-Liquidambarpollenites zone(late Middle Miocene–Late Miocene)

This zone is still dominated by angiospermouspollen (68.1–81.4%), followed by pteridophyte spores(12–32%) and gymnospermous pollen (0–9%). Themain elements in the angiospermous pollen includeCupuliferoipollenites, Quercoidites (Q. microhenrici),Ilexpollenites, Dicolpopollis, Liquidambarpollenites,JMClapapfldS

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(7–50%), and gymnospermous pollen (0–5%). Amongthe angiospermous pollen, Cupuliferoipollenites (C. ovi-formis) is predominant (generally 25%, 40% at the high-est); the herbaceous pollen component, reaching as highas 25%, includes Chenopodipollis, Echitricolporites,Artemisiaepollenites; while other commonly occur-ring types include Myricipites, Liquidambarpollenites,Saururus, Piper, Casuarinidites, Formosia, Ericipitesand Florschuetzia. The pteridophyte spores are wellrepresented by monolete types. There are only spo-radic occurrences of Pinuspollenites, Dacrydiumites,Podocarpidites and Tsugaepollenites in the gymnosper-mous pollen (Sun et al., 1981). This zone is based onthe pollen record from the Wanglougang Formation inthe Beiduwan depression and the Leiqiong areas of theNorth Continental Shelf of the South China Sea.

Pollen assemblages from the upper part of the Yue-hai Formation in the Zhujiangkou depression, the upperpart of the Yinggehai Formation in the Yinggehai depres-sion of the North Continental Shelf of the South ChinaSea (Sun et al., 1981), and the Fotan Group fromMingxi County, Fujian Province (Zheng, 1987) mightalso belong to this zone.

4.3. Inland China palynofloristic region

4.3.1. Piceaepollenites-Ulmipollenites-Chenopodipollis zone (Early Miocene)

Average values for angiospermous pollen, gym-nospermous pollen and pteridophyte spores in this zone

uglanspollenites, Alnipollenites, Ulmipollenites,yricipites and Florschuetzia, with Anodendron,henopodipollis, Ericipites and Umbelliferae having

ower values. Polypodiisporites, Polypodiaceaesporitesnd a few trilete spores dominate the pteridophyte com-onent. Only sporadic Dacrydiumites, Tsugaepollenitesnd Pinuspollenites were found in the gymnospermousollen (Sun et al., 1981). The zone is based on the pollenora from the Dengloujiao Formation of the Beiduwanepression and Leiqiong areas on the North Continentalhelf of the South China Sea.

Pollen assemblages from the upper part of the Han-iang Formation in the Zhujiangkou depression, and theower part of the Yinggehai Formation in the Yinggehaiepression on the North Continental Shelf of the Southhina Sea (Sun et al., 1981) are also comparable to thisone.

.2.4. Cupuliferoipollenites-Polypodiaceaesporites-henopodipollis zone (Pliocene)

This zone is well-represented by angiospermousollen (48–91%), followed by pteridophyte spores

are 60, 36 and 4%, respectively. The angiospermouspollen are characterized by Ulmipollenites, other amen-tiferous plants, Meliaceoidites and Engelhardtioidites.Chenopodiaceae is highly representated in the herbs,along with Leguminosae and Asteraceae. The gym-nospermous pollen component mainly contains Piceae-pollenites (4–27%), Pinuspollenites (0–18%), Tsugae-pollenites and Podocarpidites (Wang et al., 1990). Thepollen flora of this zone is reported from the upper part ofthe Xiejia Formation in the Xining-Minhe Basin, Qing-hai Province.

The palynological assemblage from the middle andupper parts of the Jidike Formation in the Kuche Basinmight also be assigned to this zone. It is marked by ahigh diversity of gymnospermous taxa including Piceae-pollenites, Pinuspollenites, Cedripites and Podocar-pidites; high representation of amentiferous plants intypes and values; a small number of tropical andsubtropical elements, such as Palmae and Trochoden-draceae (Zhao et al., 1982; Wang et al., 1990); and thePinaceae-Chenopodipollis-Betulaceae assemblage fromthe lower part of the lower Youshashan Formation


in the Qaidam Basin, Qinghai Province (Zhu et al.,1985).

4.3.2. Potamogetonacidites-Chenopodipollis-Pinaceae zone (Middle Miocene)

This zone is recorded from the Chetougou Formationin the Xining-Minhe Basin, Qinghai Province. Averagevalues for angiospermous pollen, gymnospermouspollen and pteridophyte spores are 61, 37 and 2%,respectively. Among the angiospermous pollen, bothUlmipollenites and other amentiferous plants havedeclined while herbaceous pollen has increased greatly.Increased values are recorded for Potamogetonacidites(average 17%), Chenopodipollis (average 30%),Asteraceae and Labiatae. The tropical and subtropicalelements which occurred in the previous zone havealmost disappeared. Gymnosperms are dominated byPiceaepollenites (9–47%) and Pinuspollenites (0–5%)(Sun et al., 1984; Wang et al., 1990).

The Chenopodipollis-Ephedripites-Tsugaepollenitesassemblage from the upper part of the lower YoushashanFormation in the Qaidam Basin, Qinghai Province,might also be assigned to this zone. It is distinguishedby significant values for Ephedripites (17–18%),Chenopodipollis (30%), Potamogetonacidites (18%)and Sparganiaceaepollenites (2%), and notable amountsof gymnospermous pollen including Piceaepollenites(4.5%), Abietineaepollenites (2%), Pinuspollenites(2%), and slightly higher values of Tsugaepollenites,along with the presence of Abiespollenites, Keteleeriae-

nospermous pollen. There is a very high representationof Chenopodipollis (average 56.5%) within the Neogenepollen zones in the region, and this is accompaniedby Ulmipollenites (average 12.5%), Quercoidites(average 13.5%), and Betulaceoipollenites (average4%) in the angiospermous pollen. Gymnosperms arestill represented by Pinuspollenites and Piceaepol-lenites (Sun et al., 1984; Wang et al., 1990). Thiszone is based on the pollen flora from the Xian-shuihe Formation in the Xining-Minhe Basin, QinghaiProvince.

The Pinaceae-Asteraceae-Chenopodipollis assem-blage from the upper Youshashan Formation in theQaidam Basin, Qinghai Province is characterized byhigh values and diversity of Pinaceae and over 10%Ephedripites. Herbaceous pollen rise to approach40% of total pollen, and include Asteraceae (ca. 6%),Chenopodipollis (ca. 20%), Caryophyllidites (2%),Sparganiaceaepollenites (3.5%) and Potamogetonaci-dites (ca. 10%). Woody angiospermous pollen arerather limited, and mainly represented by Nitrariadites(4–5%) and Meliaceoidites (2%) (Zhu et al., 1985). Theage of this assemblage is comparable with the zone.This zone might also include the Chenopodipollis-Artemisiaepollenites-Quercoidites-Ephedripites andChenopodipollis-Artemisiaepollenites-Ephedripites as-semblages (Ma, 1991) from the Southern DuanhuangBasin, Gansu Province.

4.3.4. Chenopodipollis-Artemisiaepollenites zone
pollenites and Cedripites (Zhu et al., 1985). This zonealso includes the Chenopodipollis-Artemisiaepollenites-Ulmipollenites-Taxodiaceaepollenites assemblage fromthe Southern Duanhuang Basin, Gansu Province (Ma,1991).
In the Inner Mongolia Subregion, the pollen florafrom the Tongguer Formation, Inner Mongolia, is domi-nated by angiospermous pollen (over 90%). Abouthalf of these represent herbaceous taxa. Themain elements are Quercoidites, Salixipollenites,Cyrillaceaepollenites, Ulmoideipites, Betulaceoipol-lenites, Rutaceoipollenites in the woody plants, andChenopodipollis, Artemisiaepollenites, Graminidites,and Compositoipollenites (syn. Echitricolporites) in theherbs. Both gymnospermous pollen and pteridophytespores have low values (Wang, 1990). This assemblagewould appear to belong to the latter part of the zone.

4.3.3. Chenopodipollis-Quercoidites-Ulmipollenites zone (Late Miocene)

This zone is dominated by angiospermous pollen(average 95%), along with a small amount of gym-

(Pliocene)Average values for angiosperm pollen, gymnosperm

pollen and pteridophyte spores in the zone are 91, 7.5and 1.5%, respectively. Herbaceous pollen, representedlargely by Chenopodipollis and Artemisiaepollenites, isthe main component. Typha, Graminidites, Echitricol-porites and Sparganiaceaepollenites are also consistentcontributors. The woody angiosperms all occur inlow values, and include Quercoidites, Salixipollenites,Liquidambarpollenites, Oleoidearumpollenites andUlmoideipites, in particular. Abietineaepollenites,Pinuspollenites, Piceaepollenites, Abiespollenites,Podocarpidites, Taxodiaceaepollenites and Ephedrip-ites are components of gymnosperm pollen, andPolypodiaceaesporites and Osmandacidites are the bestrepresented pteridophyte spores (Wang and Zhang,1990). The zone is recorded from the Shangdou-HuadeBasin in the Inner Mongolia subregion.

The Artemisiaepollenites-Ephedripites-Chenopodi-pollis assemblage (Zhu et al., 1985) reported fromthe Shizigou Formation in the Qaidam Basin, Qing-hai Province might be largely comparable with this


zone. Some pollen assemblages in the previous zoneof the Northwest China Subregion might partly extendinto the present zone. Additionally, the pollen assem-blage from the Shulehe Formation (Song, 1958) of theJiuquan Basin, Gansu Province also possibly belongshere.

4.4. Southwest China palynofloristic region

This region includes the Qinghai-Xizang and Heng-duan Mts subregions before the Middle Miocene. Sub-sequently, the subregions gradually developed inde-pendently, and formed two separate palynofloristicregions in the Pliocene. Pollen zones are based on thepollen floras from the Qinghai-Xizang area, while thosefrom the Hengduan Mts area are generally introducedas well.

4.4.1. Quercoidites-Betulaceae-Pinuspolleniteszone (late Early Miocene-early Middle Miocene)

This zone is recorded in the lower part of theWulong Formation at Namling in Xizang. It mainlycontains angiospermous pollen (generally 50–60%),along with somewhat equal occurrence of gymnosper-mous pollen and pteridophyte spores (about 10–30%).There are Quercoidites (20–30%) including Q. henrici,Q. microhenrici, Q. densus and Q. asper, Betulaceae(2–10%) containing Betulaceoipollenites, Ostryoipol-lenites and Momipites in particular and some otherthermophilic elements, such as Caryapollenites, Tiliae-piqcip1

QdaPaapPtpbmla

4.4.2. Quercoidites-Pinus-Rhododendron zone (lateMiddle Miocene)

Recovered from the upper part of the WulongFormation at Namling of Xizang, this zone is composedof 35.7–61.0% angiospermous pollen, 9.3–50.7%gymnospermous pollen, and generally 14.0–29.1%pteridophyte spores. The angiospermous pollen are stilldominated by Quercoidites, but there are higher valuesof the herbaceous taxa Ranunculaceae and Chenopo-diaceae, that exist together with Liliaceae, Gramineae,and Rhododendron. There is a higher representation ofPiceaepollenites and Abiespollenites in addition to highvalues for Pinus in the gymnospermous pollen compo-nent. Pteridophyte spores are still represented by Poly-podiisporites and Polypodiaceaesporites (Song and Liu,1982).

The pollen flora from the middle part of the Punc-tatosporites subassemblage in the Jinggu Basin, Yunnan(Song and Zhong, 1984) may be synchronous with thiszone.

4.4.3. Quercus-Salix zone (Late Miocene-earliestPliocene)

This zone is dominated by angiospermous pollen(88–91%), and contains small amounts of gymnosper-mous pollen, and sporadic pteridophyte spores. Theangiospermous pollen are mainly Quercus and Salix,that exist together with Rhus, Juglans, Betula, Celtis,Magnolia, Acer and Corylus, among the woodyplants, and Chenopodiaceae, Plantaginaceace, Liliaceae,

ollenites, Engelhardtioipollenites, Pterocaryapollen-tes, and Cupuliferoipollenites. Pinus (3–18%) is a fre-uently occurring element in the gymnosperm pollenomponent along with some Podocarpidites and Cedrip-tes. The pteridophyte spores mainly consist of Poly-odiisporites and Polypodiaceaesporites (Song and Liu,982).

In the Hengduan Mts Subregion, the Pinus-uercoidites subassemblage from 520 to 420m ofrilling core 13 in the Jinggu Basin, Yunnan is char-cterized by Abietineaepollenites, Pinuspollenites,iceaepollenites, Quercoidites, Polypodiisporitesnd Punctatosporites (Song and Zhong, 1984). Itsge might be older than the present zone, and couldossibly be assigned to the early Early Miocene. Theunctatosporites subassemblage from 368 to 271 m in

he same core could be divided into three parts. Its lowerart is comparable to the current zone. It is characterizedy diverse Quercoidites (mainly Q. minutue and Q.icrohenrici), certain Pinaceae, including Tsugaepol-

enites, and in some places pteridophyte spores (Songnd Zhong, 1984).

Gramineae and Asteraceae, among the herbs. Pinus is themajor gymnosperm pollen type (Wang et al., 1975). Thezone is recorded from the Lunpola Formation at Lunpola,Xizang.

In the Hengduan Mts. subregion, the pollen florafrom the Xiaolongtan Basin, Yunnan Province (Wang,1996) is largely comparable with this zone. It is domi-nated by angiospermous pollen, including Quercoidites(Q. minutus, Q. microhenrici, Q. henrici and Q.asper), Cupuliferoipollenites, Cyrillaceaepollenites andIlexpollenites among the woody plants, and Artemisi-aepollenites, Cichorieacidites and Tubulifloriditesamong the herbs. Abietineaepollenites, Pinuspollenites,Piceaepollenites, Tsugaepollenites and Cedripites arenotable gymnosperms, while Polypodiaceaesporites,Polypodiisporites and Polypodiaceoisporites are themost common pteridophyte spores. Some other com-parable pollen floras might include those from theupper part of the Punctatosporites subassemblage inthe Jinggu Basin, Yunnan (Song and Zhong, 1984) andthe earliest pollen stage in the Songhua Basin, YunnanProvince (Li and Wu, 1978).


4.5. Qinghai-Xizang palynofloristic region

4.5.1. Abies-Picea-Pteris zone (Early Pliocene)This zone is recorded from the Zanda Basin, Xizang.

It is located in the river valley at an altitude of3700–3750 m above sea level. The pollen flora isdominated by woody plants (70%), mostly gymnosper-mous pollen with the majority being Abies and Picea,in association with Pinus, Cedrus, a few Tsuga grainsand Podocarpus, while the angiospermous pollen taxahave relatively low values, and are represented mainlyby Betula, Alnus, Ulmus, Quercus, Juglans and Carya.There are about 10% herbs including Asteraceae,Polygonaceae, Cruciferae and Cyperaceae, and 20%pteridophyte spores, mostly represented by Pteris, andpartly by Lycopodium and Adiantum (Li and Liang,1983).

The palynological assemblage from the Jintang andCaiduo Formations at Namling, Xizang (Song and Liu,1982) might also be included in this zone.

4.5.2. Pinaceae-herbaceous pollen zone (LatePliocene)

Represented in stratigraphical beds over 3750 mabove sea level in the Zanda Basin, Xizang, this zonecan be divided into three stages. The first stage containsmainly gymnosperm pollen including Pinus and Picea(50%) and Abies (15%), a diversity of herbaceous taxawith Artemisia, Chenopodiaceae and Cyperaceae mostprominant, and a small number of other broad-leaved

monolete spores, a lower proportion of gymnosper-mous pollen (5–15%), and frequently occurring Quer-coidites, along with Betulaceae and many herbaceouspollen types (Song, 1988). Among other possibly com-parable sequences, there are the angiospermous pollensubassemblage from the Jinggu Basin (Song and Zhong,1984), the second pollen stage in the Songhua Basin (Liand Wu, 1978), and the first pollen zone from the Qujingbasin, Yunnan (Wang and Shu, 2004).

4.6.2. Pinaceae-Quercoidites pollen zone (LatePliocene)

Pollen zones 2 and 3 from the Qujing basin, Yun-nan Province are dominated by gymnospermous pollen(42.4–75.8%) including Pinus, Tsuga, Picea, Abies andCedrus, and angiospermous pollen mainly representedby Quercus in zone 2; and by angiospermous pollen(46.5–85.8%) with a majority of Quercus in zone 3(Wang and Shu, 2004). In the Songhua Basin, YunnanProvince, there are more Picea and Abies, and a morediversified herbaceous pollen flora in the upper part ofthe third stage (Li and Wu, 1978), which might be partlysynchronous with this zone.

Acknowledgements

The author thanks Peter Kershaw (Monash Uni-versity, Australia), Zhukun Zhou (Kunming Instituteof Botany, China), David K. Ferguson (University of

types, such as Salix, Ulmus, Fraxinus, and pteridophytespores. Gymnosperms are slightly lower in the secondstage, and represented by Picea and Abies, together withCedrus and Podocarpus, while the broad-leaved typesincrease, well represented by Quercus. The herbaceouspollen component is diverse and includes Chenopodi-aceae, Asteraceae, Cruciferae, Caryophyllaceae, Labi-atae, Polygonaceae, Gramineae, and many others. Thepteridophyte spores include Pteris, Onychium andLepisorus. The pollen flora becomes depauperate in thethird stage. The gymnosperms are mainly representedby Pinus, followed by Picea and Abies. The broad-leaved trees have low values except for Betula. Herba-ceous pollen is well represented by Chenopodiaceae andArtemisia (Li and Liang, 1983).

4.6. Hengduan Mts palynofloristic region

4.6.1. monolete spores-Quercoidites-herbaceouspollen zone (Early Pliocene)

Pollen zone II from the Zhaotong area, YunnanProvince, is characterized by high values for psilate

Vienna, Austria), Toshimasa Tanai (Tokyo, Japan), andTakeshi Saito (Meijo University, Japan) for their con-structive comments on the manuscript. This projectis supported by the Major Basic Research Projectsof the Ministry of Science and Technology, China(G2000077700) and the National Science Foundation ofChina (39930020).

Appendix A. Palynomorphs mentioned in thetext and their affinties

Palynomorphs Affinity

Abiespollenites Abies (Pinaceae)Abietineaepollenites Pinus (Pinaceae)Alnipollenites Alnus (Betulaceae)Artemisiaepollenites Artemisia (Asteraceae)Betulaceoipollenites BetulaceaeBetulaepollenites BetulaceaeCaryapollenites Carya (Juglandaceae)Casuarinidites CasuarinaceaeCedripites Cedrus (Pinaceae)Chenopodiaceaepollenites ChenopodiaceaeCichorieacidites Cichorieae (Asteraceae)


Appendix A (Continued )

Palynomorphs Affinity

Compositoipollenites Astereae (Asteraceae)Cupuliferoipollenites Castaneae (Fagaceae)Cyrillaceaepollenites Castaneae (Fagaceae)Dacrydiumites Dacrydium (Podocarpaceae)Deltoidospora Spore with uncertain affinityDicolpopollis PalmaeEchitricolporites Astereae (Asteraceae)Engelhardtioidites Engelhardtia (Juglandaceae)Ephedripites Ephedra (Ephedraceae)Ericipites EricaceaeFaguspollenites Fagus (Fagaceae)Florschuetzia SonneratiaceaeFonertrites Astereae (Asteraceae)Fupingopollenites Pollen with uncertain affinityGraminidites GramineaeIlexpollenites Ilex (Aquifoliaceae)Inaperturopollenites Taxodiaceae or CupressaceaeJuglanspollenites Juglans (Juglandaceae)Keteleeriaepollenites Keteleeria (Pinaceae)Laricoidites Larix (Pinaceae)Liquidambarpollenites Liquidambar (Hamamelidaceae)Lonicerapollis Lonicera (Caprifoliaceae)Lycopodiumsporites Lycopodium (Lycopodiaceae)Magnastriatites ParkeriaceaeMagnolipollis MagnoliaceaeMeliaceoidites MeliaceaeMomipites JuglandaceaeMyricipites Myrica (Myricaceae)Myrtaceidites MyrtaceaeNitrariadites Nitraria (Zygophyllaceae)Nyssapollenites Nyssa (Nyssacese)Oleoidearumpollenites OleaceaeOsmandacidites Osmunda (Osmundaceae)Ostryoipollenites Ostrya (Corylaceae)Persicariopollis Polygonaceae

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correlation of pollen sequences in the neogene palynofloristic regions of china

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