late pliocene vegetation and climate of zhangcun region...

Late Pliocene vegetation and climate of Zhangcun region,Shanxi, North ChinaF E N G Q I N *w , D AV I D K . F E R G U S O N z, R E I N H A R D Z E T T E R z, Y U F E I WA N G *, S V E T L A N A

S YA B R YA J § , J I N F E N G L I *, J I A N YA N G * and C H E N G S E N L I *

*State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Xiangshan

Nanxincun 20, Beijing 100093, China, wGraduate University of Chinese Academy of Sciences, Beijing 100049, China, zDepartment

of Palaeontology, University of Vienna, Althanstrasse 14, Vienna A-1090, Austria, §Institute of Geological Sciences, National

Academy of Sciences of Ukraine, O. Gonchara street 55-b, Kiev 01601, Ukraine

Abstract

To understand the Neogene climatic changes in eastern Asia and evaluate the intercontinental climatic differences, we

have quantitatively reconstructed the vegetation successions and climatic changes in the late Pliocene Zhangcun area

based on the palynological data and explored the regional climatic differences between central Europe and eastern

Asia. The late Pliocene palynological assemblage of Zhangcun, Shanxi was composed of 63 palynomorphs, belonging

to 50 families, covering angiosperms (90.2%), gymnosperms (9.7%), ferns (0.09%), and other elements (0.02%). Four

periods of vegetation succession over time were recognized. In period 1, a needle- and broad-leaved mixed forest

prevailed with a cool and dry climate. Period 2 was characterized by an expansion of forest with a warmer and wetter

climate. The number of conifers increased and that of herbs decreased in period 3, and the climate became cool and

dry. In period 4, the forest was dominated by conifers and reflecting a cooler climate. The data of seven climatic

parameters in general and four periods estimated by the Coexistence Approach suggested that (1) The late Pliocene

temperatures and precipitations were higher than today. (2) The Neogene climate of both Central Europe and North

China exhibited a general cooling and drying trend although the mean annual temperature dropped by ca. 1 1C in

North China, vs. ca. 7 1C in Central Europe from the middle Miocene to the late Pliocene. (3) The decline of the mean

maximum monthly precipitation might signal a weakening of the summer monsoon. (4) The decline of both the mean

coldest monthly temperature and the mean minimum monthly precipitation might be linked to the strengthening of

the winter monsoon in eastern Asia. (5) The rapid uplift of the Tibetan Plateau strengthened the climatic cooling and

drying during the late Pliocene of the Zhangcun region.

Keywords: climate, eastern Asian monsoon, late Pliocene, north China, vegetation

Received 12 August 2010; revised version received 17 November 2010 and accepted 23 November 2010

Introduction

The Neogene global climatic changes exhibited a gen-

eral cooling trend with a series of warming and cooling

fluctuations (e.g. Zachos et al., 2001; Fortelius et al., 2002;

Mosbrugger et al., 2005; Costeur & Legendre, 2008;

Utescher et al., 2009; Preto et al., 2010). Among the

terrestrial ecosystems, widespread grass-dominated

ecosystems occurred since the early to middle Miocene

(Jacobs et al., 1999; Retallack, 2001; Stromberg et al.,

2007) and C4 grassland expanded dramatically since

the late Miocene (Quade et al., 1989; Cerling et al., 1997;

Osborne, 2008; Edwards et al., 2010). More open-habitat-

adapted mammals such as grazing ungulates occurred

with the decrease of closed-forest types (Janis, 1993).

The regional differences of Neogene climatic changes

between Europe (Utescher et al., 2000; Mosbrugger et al.,

2005; Syabryaj et al., 2007; Uhl et al., 2007; Utescher et al.,

2009) and North America (Wolfe, 1994, 1995; Retallack,

2007) were revealed using different methods, e.g.

CLAMP, CA. A series of similar quantitative researches

on the Cenozoic climate in China have been accumulat-

ing since the 2000s (Sun et al., 2002; Yang et al., 2002,

2007; Liang et al., 2003; Zhao et al., 2004a; Kou et al.,

2006; Xu et al., 2008; Li et al., 2009, 2010; Xia et al., 2009;

Yao et al., 2009; Hao et al., 2010). The fossil plants found

in the late Pliocene Zhangcun Lake, Yushe Basin on the

eastern edge of Chinese Loess Plateau, provide us with

a new chance to interpret the past climatic changes in

North China.

As early as the 1930s, Teilhard & Young, (1933)

recorded the fossil mammals in the fluvial and lacus-

trine sediments of the Yushe Basin, Shanxi Province,

North China, and suggested the age as Pliocene–

Correspondence: Yufei Wang, tel. 1 86 10 6283 6439, fax 1 86 10

6259 3385, e-mail: [email protected]; Chengsen Li, tel. 1 86 10

6283 6436, fax 1 86 10 6259 3385, e-mail: [email protected]

Global Change Biology (2011) 17, 1850–1870, doi: 10.1111/j.1365-2486.2010.02381.x

1850 r 2011 Blackwell Publishing Ltd

mailto:[email protected]

mailto:[email protected]

Pleistocene. Chaney (1933) described plant megafossils

of Picea, Cyperacites, Ulmus, Ribes, Amelanchier, Legumi-

nosites and Acer from the Pliocene sediments of the

Taigu Basin. Cao & Cui (1989) reported plant mega-

fossils from the Pliocene Zhangcun Formation of the

Yushe Basin, including Picea, Pinus, Salix, Populus,

Juglans, Pterocarya, Carpinus, Quercus, Ulmus, Zelkova,

Hemiptelea, Cudrania, Broussonetia, Spiraea, Albizia, Acer,

Euodia, Koelreuteria, Typha, Potamogeton and Zannichellia,

suggesting a warm temperate deciduous broad-leaved

forest and a semihumid warm temperate monsoon

climate. Later, Zhao et al. (2004b) recorded Ruppia

yushensis from the Zhangcun Formation, suggesting

that it lived in a brackish, clear, tranquil and shallow

lake under a warm temperate or temperate climate. Liu

et al. (2005) described Bolboschoenus cf. yagara from the

Zhangcun Formation, implying a wetland environment.

Shi et al. (1993) reported the pollen assemblages from

the Yushe Group of Yushe Basin belonging to the

Pliocene to early Pleistocene, including Rejianao For-

mation, Wangning Formation, Zhangcun Formation

and Louzeyu Formation from bottom to top. They

suggested that a cool and dry climate prevailed in the

middle and later period of the Zhangcun Formation,

based on the increase in pollen percentages of conifers

and xerophytes.

Liu et al. (2002) described the Pliocene (3.2–2.0 Ma)

pollen assemblages of the Yushe Basin and pointed out

that while Zone A suggested an open deciduous wood-

land with steppe under a warm temperate climate with

moist summers, Zone B indicated a coniferous forest

growing under a relatively cooler semihumid temperate

climate.

Li et al. (2004) studied the Zhangcun pollen assem-

blages (2.77–2.52 Ma) of the Zhangcun Formation and

suggested that the palaeovegetation developed from

coniferous forest, broad-leaved forest and steppe, to

coniferous forest, reflecting fluctuations in the climate

from cold-wet, relatively warm-dry to cold-wet.

In this article, we reconstruct the vegetation succes-

sions and climatic changes in the late Pliocene Zhang-

cun area by the Coexistence Approach (CA) based on

new palynological data, and explore the regional cli-

matic differences between central Europe and eastern

Asia.

Materials and methods

Study site

The Zhangcun site, Yushe Basin of Shanxi Province is located

on the eastern edge of the Chinese Loess Plateau, and falls in

the modern Chinese warm temperate monsoon zone (CCPGC,

1984).

The Yushe Basin, a dendritic saucer-like synclinal basin, is

located in middle-east Shanxi extending in a NNE–SSW direc-

tion. The late Cenozoic fluvial and lacustrine sediments un-

conformably overlie the Triassic arenaceous shale in the Yushe

Basin (Cao & Wu, 1985).

Huang & Guo (1991) summarized the history of strati-

graphic subdivision of the late Cenozoic sediments of the

Yushe Basin. Shi et al. (1993) dated these late Cenozoic strata

as ca. 5.5–1.5 Ma using palaeomagnetism, and divided them

into four formations from bottom to top, i.e. Renjianao Forma-

tion (5.5–4.6 Ma), Wangning Formation (4.6–3.5 Ma), Zhangcun

Formation (3.5–2.3 Ma) and Luozeyu Formation (2.3–1.5 Ma).

The Zhangcun section (361580N, 1121510E, 1043 m asl,

15.66 m thick, Figs 1 and 2) is in the middle of the upper part

of the Zhangcun Formation and is dated from 2.77 to 2.52 Ma

(Li et al., 2004). The study profile from bottom to top is divided

into 13 layers. The lithostratigraphic characters of each layer

are described as follows:

Overlying strata: Quaternary loess

Unconformity

13 Gray-yellow muddy diatomite 2.00 m

12 Gray muddy diatomite 1.90 m

11 Gray-white muddy diatomite 5.60 m

10 Light gray-yellow diatomaceous shale 0.40 m

9 Gray-white diatomaceous shale 1.00 m

8 Light gray-yellow diatomite 0.20 m

7 Gray-white muddy diatomite 0.70 m

6 Gray-white muddy diatomite with 9

intercalated layers of gypsum 0.2–1 cm

thick at 4–9 cm intervals

0.86 m

5 Light gray diatomite with 14 intercalated

layers of gypsum 0.2–1 cm thick

at 1–6 cm intervals

0.55 m

4 Dark gray diatomite with 3 intercalated

layers of gypsum 0.5–1 cm thick at

8–22 cm intervals, yellow siltstone of

10–15 cm thick in the upper and

lower parts

0.60 m

3 Gray clay with 4 intercalated layers of

gypsum 1–2 cm thick at 6–11 cm intervals

0.65 m

2 Dark gray clay 0.70 m

1 Gray clay 0.50 m

Conformity

Underlying strata: yellow siltstone

Materials and methods

Eighty-eight palynological samples were collected from the

Zhangcun section (Fig. 3). In the portion containing interca-

lated layers of gypsum, samples were collected in, above and

below the gypsum.

The samples were treated by the method of Heavy Liquid

Separation (density: 2.0 g mL�1; Moore et al., 1991; Li & Du,

L A T E P L I O C E N E V E G E TA T I O N A N D C L I M A T E 1851

r 2011 Blackwell Publishing Ltd, Global Change Biology, 17, 1850–1870

1999). The single-grain technique of Ferguson et al. (2007) was

applied. The pollen grains and spores were observed under a

scanning electron microscope (SEM). Usually more than 200

pollen/spore grains within one or two slides per sample are

counted in the pool of samples, by using a Leica DM 2500 light

microscope (LM). Pollen grains and spores were found in all

Yushe

Zhangcun

37°00′ 37°00′

112°55′

112°55′

4km N

113°00′

113°00′

112°50′

112°50′

37°05′37°05′

36°55′ 36°55′

Zhu

ozha

ngR

iver

Fig. 1 Maps showing the position of the Zhangcun locality.

Fig. 2 Zhangcun sampling locality. (a) A general view of the Zhangcun locality. (b) An outcrop of the middle-upper part of the

Zhangcun Formation. (c) Gypsum layers (arrows) in the profile. (d) Gypsum crystals. (e) A grasslike leaf fragment collected from the

profile. Scale bar 5 1 cm.

1852 F. Q I N et al.


samples, fewer than 100 grains (3–83) were found in 10

samples (samples number zc18, 65, 67, 77, 82, 83, 84, 85, 86

and 87), while more than 100 grains (116–5423) were found in

the other 78 samples. Those samples with fewer than 100

pollen/spore grains (3–83) within three slides are not plotted

in the distribution of relative abundances of main taxa in the

measured Zhangcun section (Fig. 4) for the number of those

pollen/spores preserved in the samples is too few to reflect the

local vegetation composition at that time objectively.

The palynomorph relative abundance (RA) of a taxon is

calculated by the equation: RA 5 N/Nt, where N is the the

pollen/spore number of a taxon and Nt represents the total

Zc1Zc2Zc3Zc4Zc5Zc6

Zc88

Zc87

Zc84

Zc86Zc85

Zc83

Zc82

Zc81

Zc80

Zc79

Zc78

Zc77Zc76Zc75Zc74Zc73Zc72Zc71

SiltstoneClay

DiatomiteMuddy diatomiteLoess

Gypsum layer

Zc70Zc69Zc68Zc67Zc66Zc65Zc64Zc63Zc62 Zc61

Zc60

Zc52

Zc59Zc58Zc57Zc56Zc55Zc54Zc53Zc51Zc50

Zc49Zc48Zc47Zc46Zc45

Zc38Zc37Zc36Zc34Zc33Zc32Zc31Zc30Zc29Zc28Zc27

Zc25

Zc44Zc43Zc42Zc41Zc40 Zc35

Zc39

Zc24Zc23Zc22Zc21Zc20Zc19

Zc18

Zc17Zc16Zc15Zc14Zc13Zc12Zc11Zc10Zc9

Zc8Zc7

Zc26

Mudstone

Diatomaceousshale

Quaternarysediment

0 00m.

Layer 13

Layer 12

Layer 11

Layer 10

Layer 9Layer 8Layer 7

Layer 6

Layer 5Layer 4

Layer 3

Layer 2Layer 1

Layer 6

Layer 5

Layer 4

Layer 3

2 00m.

3 90m.

9 50m.9 90m.

10 90m.11 10m.

11 80m.

12 66m.13 21m.13 81m.

14 46m.

15 16m.15 66m.

Yellowsiltstone

Zone I

Zone II

Zone III

Zone IV

Fig. 3 Measured stratigraphical sequence of Zhangcun.



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pollen/spore number of all taxa combined in the pool of

samples. TILIA software and TILIAGRAPH was used to

construct the pollen diagram ( 5 the distribution of relative

abundances of main taxa in the measured Zhangcun section,

Fig. 4). The pollen and spores were identified by referring to

the palynological literature and monographs (IBCAS, 1976;

IBCAS & SCIBCAS, 1982; Xi & Ning, 1994; Wang et al., 1995).

The late Pliocene climate of Zhangcun was reconstructed

following the CA (Mosbrugger & Utescher, 1997). Seven para-

meters of temperature and precipitation were estimated, i.e.

the mean annual temperature (MAT), the mean warmest

monthly temperature (MWMT), the mean coldest monthly

temperature (MCMT), the temperature difference between

coldest and warmest months (DT), the mean annual precipita-

tion (MAP), the mean maximum monthly precipitation

(MMaP), and the mean minimum monthly precipitation

(MMiP). To estimate the range of climatic variables, we inte-

grated the geographical distributions of the nearest living

relatives (NLRs) of pollen taxa in China (Wu & Ding, 1999)

and the modern surface meteorological data (1951–1980) with-

in their distribution areas (IDBMC, 1983a, b, c, 1984a, b, c). In

addition, the MAT values of NLRs from the Palaeoflora

Database were also adopted (http://www.geologie.uni-

bonn.de/Palaeoflora/Palaeoflora_home.htm).

Results

The Zhangcun palynoflora (Table 1, Plate I–V) yielded

63 palynomorphs assigned to 50 families, covering

angiosperms (40 families), gymnosperms (three fa-

milies), pteridophytes (five families) and algae (two

families). The relative abundance of angiosperm pollen

in this palynoflora was 90.2%, among which herbs

(Artemisia 66.2% and Chenopodiaceae 10.8%) composed

78.5%, trees 11.7% (Ulmus/Zelkova 7.5%, Corylus 1.8%,

Table 1 The palynomorph relative abundance (RA) of

Zhangcun section

Palynomorph RA (%)

Algae 0.02

Pediastrum 0.01

Zygnemataceae 0.01

Pteridophytes 0.09

Polypodiaceae 0.05

Athyriaceae 0.01

Hemionitidaceae 0.01

Pteridaceae 0.01

Selaginellaceae 0.01

Gymnosperms 9.69

Pinus 4.74

Abies 3.47

Picea 0.81

Larix 0.42

Ephedra 0.19

Taxodiaceae 0.05

Tsuga 0.02

Angiosperms 90.19

Artemisia 66.20

Chenopodiaceae 10.79

Ulmus/Zelkova 7.46

Corylus 1.80

Quercus 0.78

Betula 0.54

Eucommia 0.44

Brassicaceae/Cruciferae 0.37

Ranunculaceae 0.30

Juglans 0.24

Asteraceae/Compositae 0.23

Typha 0.18

Tilia 0.13

Poaceae/Gramineae 0.12

Fabaceae/Leguminosae 0.11

Potamogeton 0.09

Alnus 0.07

Carya 0.06

Polygonum 0.06

Oleaceae 0.04

Sparganium 0.03

Cyperaceae 0.02

Lamiaceae/Labiatae 0.02

Anacardiaceae 0.01

Campanulaceae 0.01

Caprifoliaceae 0.01

Castanea 0.01

Elaeagnus 0.01

Fraxinus 0.01

Rosaceae 0.01

Apiaceae/Umbelliferae 0.01

Acer /

Araliaceae /

Caryophyllaceae /

Convolvulaceae /

Continued

Table 1. (Contd.)

Palynomorph RA (%)

Dipsacaceae /

Euphorbiaceae /

Erodium /

Ilex /

Malvaceae /

Nitraria /

Onagraceae /

Oxalidaceae /

Populus /

Pterocarya /

Rutaceae /

Salix /

Viola /

Zygophyllaceae /

/, the relative abundance of the palynomorph is below 0.01%.



http://www.geologie.uni-bonn.de/Palaeoflora/Palaeoflora_home.htm

http://www.geologie.uni-bonn.de/Palaeoflora/Palaeoflora_home.htm

Plate I 1–3, Taxodiaceae; 4–7, Abies; 8–11, Picea; 12–14, Larix; 15–17, Acer. Scale bar in light microscope (LM) and scanning electron

microscopic (SEM) overview 10 mm, in SEM close-up 1mm.



Plate II 18–23, Quercus; 24–26, Oleaceae; 27–29, Araliaceae; 30–32, Salix. Scale bar in light microscope (LM) and scanning electron




Plate III 33–35, Ulmus; 36–38, Zelkova; 39–41, Alnus; 42–44, Betulaceae; 45–47, Corylus. Scale bar in light microscope (LM) and scanning

electron microscopic (SEM) overview 10 mm, in SEM close-up 1mm.



Plate IV 48–53, Eucommia; 54–56, Elaeagnus; 57–59, Juglans; 60–62, Carya. Scale bar in light microscope (LM) and scanning electron




Plate V 63–65, Artemisia; 66–68, Chenopodiaceae; 69–71, Poaceae/Gramineae; 72–74, Typha; 75–77, Athyriaceae. Scale bar in light

microscope (LM) and scanning electron microscopic (SEM) overview 10 mm, in SEM close-up 1mm.



Quercus 0.8%, Betula 0.5% and Eucommia 0.4%); gym-

nosperms 9.7% (Pinus 4.7%, Abies 3.5% and Picea 0.8%);

pteridophytes 0.09% and algae 0.02%.

The Zhangcun palynomorph taxa (Table 2) were

grouped by the temperature preferences of their nearest

living relatives (Jimenez-Moreno, 2006; Li et al., 2009) as

one megathermic element, two mega-mesothermic ele-

ments, 17 mesothermic elements, two mesomicrother-

mic elements, three microthermic elements, six

nonsignificant element and 24 herbs and/or shrubs.

Based on the palynomorphs and their relative abun-

dances, the pollen spectra (Fig. 4) were divided into

four pollen zones.

Zone I (15.7–14.0 m, 12 samples): 47 types of palyno-

morphs were identified, i.e. 36 angiosperms, six gym-

nosperms, three pteridophytes and two algae (Table 2).

The relative abundance of angiosperm pollen repre-

sented a total of 94.9%, of which herbs 92% and trees

2.9%. Of the herbs, Artemisia (63.6%) and Chenopodia-

ceae (27.7%) were dominant elements. Asteraceae/

Compositae (0.3%), Ranunculaceae (0.2%) and Brassi-

caceae/Cruciferae (0.1%) were also found. Aquatic

and/or marsh plants like Potamogeton (0.02%), Typha

(0.01%) and Sparganium (o0.01%) were rare. Among

Table 2 The list of the late Pliocene Zhangcun palynomorph

taxa grouped by ecological requirements and their relative

abundance (RA) in the four pollen zones (table style based on

Jimenez-Moreno, 2006 and Li et al., 2009)

Zones I (%) II (%) III (%) IV (%)

Megathermic elements

Rutaceae – / – –

Mega-mesothermic elements

Anacardiaceae 0.02 0.01 – –

Eucommia 0.04 0.68 – –

Taxodiaceae 0.01 0.06 0.10 2.53

Mesothermic elements

Acer – / – –

Alnus 0.01 0.11 – –

Betula 0.71 0.45 0.72 0.42

Carya 0.04 0.08 – –

Castanea 0.03 0.01 – –

Corylus 0.39 2.60 1.64 –

Elaeagnus 0.03 – – –

Fraxinus – 0.01 – –

Ilex – / – –

Juglans 0.10 0.32 0.05 –

Oleaceae 0.08 0.02 0.05 –

Populus – – – 0.42

Pterocarya / – – –

Quercus 0.32 1.04 0.62 0.42

Salix – / – –

Tilia 0.08 0.15 0.10 –

Ulmus/Zelkova 0.94 10.95 11.28 0.42

Meso-microthermic elements

Pinus 1.92 5.70 15.54 35.02

Tsuga – 0.02 0.10 0.42

Microthermic elements

Abies 2.04 3.01 29.38 38.82

Larix 0.15 0.57 0.51 –

Picea 0.74 0.63 3.79 18.57

Non-significant elements

Araliaceae / / – –

Caprifoliaceae 0.01 / – –

Euphorbiaceae – / – –

Fabaceae/Leguminosae 0.06 0.14 – –

Malvaceae – / – –

Rosaceae / 0.01 – –

Herbs and shrubs

Apiaceae/Umbelliferae 0.02 / – –

Artemisia 63.62 69.30 33.64 1.27

Asteraceae/Compositae 0.32 0.19 0.15 –

Brassicaceae/Cruciferae 0.13 0.50 0.62 –

Campanulaceae – 0.01 – –

Caryophyllaceae / / – –

Chenopodiaceae 27.67 1.91 1.23 0.84

Convolvulaceae 0.01 / – –

Cyperaceae 0.02 0.01 0.10 –

Dipsacaceae 0.01 – – –

Continued

Table 2. (Contd.)

Zones I (%) II (%) III (%) IV (%)

Ephedra 0.10 0.24 0.05 0.42

Erodium / – – –

Lamiaceae/Labiatae / 0.03 – –

Nitraria – / – –

Onagraceae – / – –

Oxalidaceae – / – –

Poaceae/Gramineae 0.04 0.17 – –

Polygonum 0.01 0.08 – –

Potamogeton 0.02 0.14 – –

Ranunculaceae 0.16 0.38 0.21 0.42

Sparganium / 0.04 – –

Typha 0.01 0.28 – –

Viola / – – –

Zygophyllaceae – / – –

Pteridophytes

Athyriaceae – 0.01 – –

Hemionitidaceae 0.01 0.01 – –

Polypodiaceae 0.09 0.03 – –

Pteridaceae – 0.01 – –

Selaginellaceae 0.01 0.01 0.10 –

Other elements

Pediastrum 0.01 0.02 – –

Zygnemataceae 0.01 0.01 – –

–, no pollen grain or spore of this taxon exists; /, the relative

abundance of the palynomorph is below 0.01%.



arboreal angiosperms, Ulmus/Zelkova (0.9%), Betula

(0.7%), Corylus (0.4%), Quercus (0.3%) and Tilia (0.1%)

were dominant. The gymnospermous pollen contribu-

ted 5%, including Abies (2%), Pinus (1.9%), Picea (0.7%),

Larix (0.2%), Ephedra (0.1%) and Taxodiaceae (0.01%).

The most common fern was Polypodiaceae with a

relative abundance of 0.1%.

No megathermic element appeared in this zone.

There were three megamesothermic elements (Anacar-

diaceae, Eucommia and Taxodiaceae), 12 mesothermic

elements (e.g. Betula, Corylus, Quercus, Ulmus/Zelkova),

one mesomicrothermic element (Pinus), three micro-

thermic elements (Abies, Larix and Picea), four nonsigni-

ficant elements (Araliaceae, Caprifoliaceae, Fabaceae/

Leguminosae and Rosaceae) and 19 kinds of herbs and/

or shrubs (e.g. Artemisia, Chenopodiaceae, Asteraceae/

Compositae, Ranunculaceae) in this zone (Table 2).

Zone II (14.0–10.0 m, 64 samples): 57 types of palyno-

morphs were found, covering 43 angiosperms, seven

gymnosperms, five pteridophytes and two algae (Table 2).

In comparison with Zone I, the relative abundance of

herbaceous angiosperm pollen (73%) decreased, Arte-

misia (69.3%) increased while Chenopodiaceae (1.9%)

decreased sharply. Arboreal angiosperms (16.6%), espe-

cially Ulmus/Zelkova (11%), Corylus (2.6%), Quercus (1%),

Eucommia (0.7%), and Alnus (0.1%) increased.

The gymnosperms (10.2%) increased, of which Pinus

contributed 5.7% and Abies 3%.

One megathermic element (Rutaceae), three megame-

sothermic elements (Anacardiaceae, Eucommia and Tax-

odiaceae), 14 mesothermic elements (e.g. Betula, Corylus,

Quercus, Ulmus/Zelkova), two mesomicrothermic ele-

ments (Pinus and Tsuga), three microthermic elements

(Abies, Larix and Picea), six nonsignificant elements (e.g.

Araliaceae, Caprifoliaceae, Euphorbiaceae, Fabaceae/

Leguminosae) and 21 kinds of herbs and/or shrubs

(e.g. Artemisia, Chenopodiaceae, Brassicaceae/Cruciferae,

Ranunculaceae) were found (Table 2).

Zone III (10.0–5.0 m, 5 samples): 21 palynomorphs were

recorded, including 13 angiosperms, seven gymnos-

perms and one pteridophyte (Table 2).

In comparison with Zone II, the relative abundance of

herbaceous angiosperm pollen (e.g. Artemisia 33.6%,

Chenopodiaceae 1.2%) and arboreal angiosperms (e.g.

Corylus 1.6% and Quercus 0.6%; Eucommia and Alnus

absent) decreased. Ulmus/Zelkova was still the most

important arboreal angiosperm with a relative abun-

dance of 11.3%. However, the pollen of gymnosperms

such as Abies (29.4%), Pinus (15.5%) and Picea (3.8%),

reached 49.5% of the total pollen counts.

There was no megathermic element, one megame-

sothermic element (Taxodiaceae), seven mesothermic

elements (e.g. Betula, Corylus, Quercus, Ulmus/Zelkova),

two meso-microthermic elements (Pinus and Tsuga),

three microthermic elements (Abies, Larix and Picea),

and seven herbs and/or shrubs (e.g. Artemisia, Brassi-

caceae/Cruciferae, Cyperaceae, Ranunculaceae) in this

zone (Table 2).

Zone IV (5.0–0 m, 7 samples): 13 types of palynomorphs

were found, belonging to seven angiosperms and six

gymnosperms (Table 2).

Coniferous pollen dominated the pollen assemblage

with a relative abundance of 95.4%, including Abies

(38.8%), Pinus (35%), Picea (18.6%), Tsuga (0.4%) and

Taxodiaceae (2.5%). The relative abundance of angios-

perm pollen was only 4.2%.

One megamesothermic element (Taxodiaceae), four

mesothermic elements (Betula, Populus, Quercus and

Ulmus/Zelkova), two mesomicrothermic elements (Pi-

nus and Tsuga), two microthermic elements (Abies and

Picea), and four herbs and/or shrubs (Artemisia, Cheno-

podiaceae, Ephedra, Ranunculaceae) were found in this

zone (Table 2).

Discussion

Qualitative analysis of the palaeovegetation andpalaeoclimate

The whole pollen assemblage of the late Pliocene of

Zhangcun suggested a mixed deciduous broad-leaved

and coniferous forest, which indicates a warm tempe-

rate climate. The NLRs of the main taxa (e.g. Abies,

Picea, Pinus, Betula, Corylus, Juglans, Quercus, Tilia,

Ulmus, Zelkova) of the forest are widespread in the

temperate zone and/or montane areas of the subtropi-

cal zone today. Thermophilous elements (e.g. Rutaceae,

Eucommia) grow sparsely in lowland. Artemisia and

Chenopodiaceae, as predominant herbs, are mainly

found in the arid and semiarid areas of the temperate

zone. The aquatic plants, Typha, Potamogeton and Spar-

ganium are distributed widely in the temperate zone.

Today it is obvious that some pollen may be trans-

ported long distances (Cour et al., 1999; Rousseau et al.,

2003, 2006, 2008). Nevertheless, we here notice that

many common and bio-sensitive plant elements are

found in both Zhangcun Pliocene palynoflora (this

paper) and mega-fossil leaf/fruit flora (Cao & Cui,

1989), e.g. Ulmus, Zelkova, Quercus, Picea, Pinus, Juglans,

Acer, Salix, Populus, Typha, Eucommia (a Eucommia fruit

was recorded in Wang, 2009). Of these, Ulmus, Zelkova,

Quercus, Picea, Pinus were no doubt dominant elements

in the palynoflora. The palynoflora could be used to

reflect the past climate conditions for it received most

pollen from numerous plants nearby although it also

included some pollen by long-distance transport.

Based on the palynomorphs and their relative abun-

dances from bottom to top, four periods of vegetation



succession in the Zhangcun region during the late

Pliocene were recognized as follows:

Period 1 (Zone I, Fig. 5): A needle- and broad-leaved

mixed forest prevailed in Zhangcun during this period.

Typical temperate elements such as Pinus, Betula,

Corylus, Juglans, Quercus, Tilia and Ulmus/Zelkova,

dominated the forest. In addition, thermophilous ele-

ments like Anacardiaceae, Araliaceae, and Eucommia

and psychrophilous ones, e.g. Abies, Picea and Larix

were present. Ferns and fern allies like Polypodiaceae,

Hemionitidaceae and Selaginellaceae grew in the her-

baceous layer of the forest.

Hygrophilous elements such as Alnus and Taxodia-

ceae grew sparsely around the palaeolake, while aqua-

tic plants like Typha, Potamogeton and Sparganium grew

in the palaeolake or surrounding wetland.

Xerophilous elements such as Chenopodiaceae and

Artemisia dominated the herbaceous vegetation on the

lakeshore. Some Ephedra also existed on the dry slopes.

The relatively higher pollen ratio (0.07–1.46) of Cheno-

podiaceae/Artemisia indicated dry conditions (El-Mos-

limany, 1990).

The area was characterized by a warm and dry

climate during this period.

Period 2 (Zone II, Fig. 5): In comparison with Period 1,

the needle- and broad-leaved mixed forest expanded

dramatically. The broad-leaved trees such as Corylus,

Quercus, Ulmus/Zelkova, and conifers such as Abies,

Pinus dominated the forest. The number and/or diver-

sity of thermophilous elements (Eucommia, Euphorbia-

ceae, Ilex and Malvaceae) and hygrophilous ones, Alnus

and Taxodiaceae increased.

Xerophilous elements like Chenopodiaceae and Arte-

misia continued to predominate in the herbaceous ve-

getation lining the lakeshore. However, the pollen ratio

of Chenopodiaceae/Artemisia (0.01–0.28) declined with

a sharp decrease of Chenopodiaceae, indicating a wetter

habitat.

The pollen assemblage indicated a warmer and wet-

ter climate during this period.

Period 3 (Zone III, Fig. 5): In comparison with Period 2,

the number of conifers such as Abies, Picea and Pinus

increased sharply. Ulmus/Zelkova remained the predo-

minant element in broad-leaved types, but the number

of Corylus and Quercus decreased. All the thermophi-

lous elements except Taxodiaceae disappeared.

The number of Artemisia decreased sharply although

it still predominated in the herb layer. The pollen ratio

HerbsAquaticplants

Shrubs Broad-leavedtrees

Abies and Picea

Period 3 Period 4

Period 1 Period 2

Pinus

Fig. 5 The vegetation succession of the Zhangcun region from Periods 1 to 4 during the late Pliocene.



of Chenopodiaceae/Artemisia (0.03–0.79) was higher at

the beginning of this period, implying drier conditions.

A cool and dry climate is indicated by the pollen

assemblage of this period.

Period 4 (Zone IV, Fig. 5): In this period, conifers such

as Abies, Picea, Pinus and Taxodiaceae dominated the

forest vegetation. The number and diversity of both

arboreal and herbaceous angiosperms decreased shar-

ply. The pollen of Artemisia, Betula, Chenopodiaceae,

Cyperaceae, Populus, Quercus, Ranunculaceae and

Ulmus/Zelkova represented o1% of the total pollen

counts.

This undoubtedly indicates a cooler climate during

period 4.

On the whole, the late Pliocene Zhangcun region

experienced a warm-dry, warm-wet, cool-dry, to cool

climate during periods 1–4.

Quantitative reconstruction of palaeoclimate by CA

Based on the whole pollen assemblage (Table 1), the

data of seven climate parameters were estimated as

follows: MAT 5 8.5–15.1 1C, delimited by Castanea and

Nitraria, MWMT 5 19.8–27.5 1C, delimited by Taxodia-

ceae and Betula, MCMT 5�0.3 to 2.0 1C, delimited by

Carya and Nitraria, DT, MAP and MMaP were 25.0–

26.0 1C, 845.6–1050.9 and 183.6–229.4 mm, respectively,

all delimited by Eucommia and Nitraria, MMiP 5 19.2–

21.2 mm, delimited by Taxodiaceae and Eucommia

(Fig. 6, Table 3).

Compared with the current meteorological data, all

the median values of MAT, MWMT, MCMT, MAP,

MMaP and MMiP except DT were higher during the

late Pliocene (Table 4), implying a warmer and wetter

climate with weaker seasonality in the late Pliocene of

Zhangcun.

Unfortunately, there are no quantitative climatic re-

sults derived from sources other than plants in this

region. Nevertheless, the occurrence of fossil mammals

like Rhinoceros, Trilophodon, Tetralophodon, and Stegodon

found in the Pliocene–Pleistocene sediments of the

Yushe Basin (Teilhard & Young, 1933; Licent & Tras-

saert, 1935) probably indicate a warmer climate than

today, e.g. the modern Rhinoceros, as the nearest living

relative of fossil Rhinoceros lives in tropical or subtropi-

cal climate conditions.

The climatic fluctuations from Zones I–IV

The palaeoclimatic fluctuations can be illustrated by the

curves of median values of seven climatic parameters

during the periods of time represented by Zones I–IV.

Here, we calculate the arithmetic mean of the upper and

lower boundary of the coexistence interval, which

is used to obtain the median value of each climatic

parameter.

MAT (Fig. 7a; Table 3): The MAT values declined from

Zone I (8.5–20.9 1C) to Zone II (8.5–15.1 1C), went up in

Zone III (7.8–22.1 1C) and were then followed by less

oscillation in Zone IV (7.8–22.7 1C).

MWMT (Fig. 7b; Table 3): The MWMT values re-

mained uniform (19.8–27.5 1C) during Zones I–IV.

MCMT (Fig. 7c; Table 3): The MCMT values fell from

Zone I (�0.3 to 5.9 1C) to Zone III (�8.8 to 5.9 1C), then

remained stable from Zone III to Zone IV (�8.8 to

5.9 1C).

DT (Fig. 7d; Table 3): The DT values increased from

Zone I (15.8–26.0 1C) to Zone II (25.0–26.0 1C), declined

from Zone II to Zone III (13.0–33.1 1C), then remained

constant from Zone III to Zone IV.

MAP and MMaP (Fig. 7e and f; Table 3): The fluctua-

tions of MAP (573.9–1254.7 mm) and MMaP (129.4–

245.2 mm) values paralleled those of MCMT.

MMiP (Fig. 7g; Table 3): The MMiP values remained

constant in both Zone I and Zone II (19.2–21.2 mm), then

oscillated between 5.7 and 21.2 mm in Zones III–IV.

Comparison with the Neogene climate of Central Europe

The Neogene climatic changes in North China may be

described by the curves of median values of climatic

parameters in the early Miocene of Weichang (Li et al.,

2009), the middle Miocene of Shanwang (Yang et al.,

2007) and the late Pliocene of Zhangcun (this paper).

In North China, the MAT curve reflected a warming

period from the early to middle Miocene, then a cooling

trend from the middle Miocene to the late Pliocene

(Table 5, Fig. 8), which coincided with the general trend

of global MAT fluctuations (e.g. Zachos et al., 2001;

Mosbrugger et al., 2005).

Compared with Central Europe, the MAT curve of

North China exhibited the same trend but different

amplitudes from the middle Miocene to late Pliocene,

i.e. MAT dropped by ca. 1 1C in North China but ca. 7 1C

in Central Europe, although the MAT values increased

by ca. 1 1C in both North China and Central Europe

from the early to middle Miocene.

The MWMT and MCMT curves of North China and

Central Europe exhibit different trends (Table 5, Fig. 8).

The MWMT values started from the early to middle

Miocene with a cooling trend in North China vs. a

warming trend in Central Europe, and then stabilized

in North China as opposed to the cooling which occurred

in Central Europe from the middle Miocene to the late

Pliocene. The MCMT values displayed a cooling trend

from the early Miocene to the late Pliocene in North

China, while an amelioration from the early to middle



105

1520253035

−10−20

0102030

−30

2010

304050

0

1200800

1600200024002800

4000

0−5

510152025

−10

1 10 20 30 40 50 57

200100

300400500600

0

DT

(°C

)M

AT

(°C

)M

WM

T(°

C)

MC

MT

(°C

)M

AP

(mm

)M

MaP

(mm

)M

MiP

(mm

)

4020

6080

019.2mm21.2mm

183.6mm229.4mm

845.6mm1050.9mm

26.0°C25.0°C

2.0°C−0.3°C

27.5°C19.8°C

15.1°C8.5°C

Fig. 6 The coexistence interval of the climatic parameters of Zhangcun palynomorphs. 1, Abies; 2, Acer; 3, Alnus; 4, Anacardiaceae; 5,

Apiaceae/Umbelliferae; 6, Araliaceae; 7, Artemisia; 8, Asteraceae/Compositae; 9, Betula; 10, Brassicaceae/Cruciferae; 11, Campanu-

laceae; 12, Caprifoliaceae; 13, Carya; 14, Caryophyllaceae; 15, Castanea; 16, Chenopodiaceae; 17, Convolvulaceae; 18, Corylus; 19,

Cyperaceae; 20, Dipsacaceae; 21, Elaeagnus; 22, Ephedra; 23, Erodium; 24, Eucommia; 25, Euphorbiaceae; 26, Fabaceae/Leguminosae; 27,

Fraxinus; 28, Ilex; 29, Juglans; 30, Lamiaceae/Labiatae; 31, Larix; 32, Malvaceae; 33, Nitraria; 34, Oleaceae; 35, Onagraceae; 36,

Oxalidaceae; 37, Picea; 38, Pinus; 39, Poaceae/Gramineae; 40, Polygonum; 41, Populus; 42, Potamogeton; 43, Pterocarya; 44, Quercus; 45,

Ranunculaceae; 46, Rosaceae; 47, Rutaceae; 48, Salix; 49, Sparganium; 50, Taxodiaceae; 51, Tilia; 52, Tsuga; 53, Typha; 54, Ulmus; 55, Viola;

56, Zelkova; 57, Zygophyllaceae.



Miocene was followed by a cooling trend during the

middle Miocene to the late Pliocene in Central Europe.

The curve of MAP increased by ca. 470 mm from the

early to middle Miocene and then dropped by ca.

550 mm from the middle Miocene to late Pliocene in

North China. However, in Central Europe, the MAP

values fluctuated mostly from 1200 to 1300 mm during

the Miocene and then dropped suddenly by ca. 250 mm

in the early Pliocene (Table 5, Fig. 8).

Palaeomonsoon

The Neogene climatic changes in North China differ

from those of Central Europe due to their unique

topography (Tapponnier et al., 2001; Spicer et al., 2003;

Royden et al., 2008) and Asian monsoons (Burbank et al.,

1993; Kutzbach et al., 1993; An et al., 2001) caused by the

rapid uplift of the Tibetan Plateau since the Miocene

(Harrison et al., 1992; An et al., 2006; Zhang et al., 2008).

The uplift of the Tibetan Plateau caused by the India-

Asia collision event since the early Eocene (Rowley,

1996) not only modified the topography (Wang & Li,

1993; Li & Fang, 1998) and the regional climate of China

(Dettman et al., 2003; Kou et al., 2006; Wang et al., 2006;

Xia et al., 2009; Yao et al., 2009), but also influenced the

global climate (Ruddiman & Raymo, 1988; Raymo &

Ruddiman, 1992; Molnar et al., 1993; Jiang et al., 2008)

and the intensity of the Asian monsoons (Prell &

Kutzbach, 1992; Kutzbach et al., 1993).

The rapid uplift of the Tibetan Plateau during the

Pliocene (Zheng et al., 2000; Li et al., 2001; An et al., 2006;

Zhang et al., 2008) profoundly influenced the climate

and monsoon intensity on the Chinese Loess Plateau

(Ding et al., 1998, 2005; Qiang et al., 2001; Wu et al., 2006;

Bai et al., 2009). An et al. (2001) pointed out that during

the late Pliocene, both eastern Asian summer and

winter monsoons intensified from ca. 3.6 to 2.6 Ma,

while the eastern Asian winter monsoon continued

strengthening with the possible weakening of the east-

ern Asian summer monsoon after ca. 2.6 Ma, based on

the records of aeolian sediments from China (Sun et al.,

1997, 1998) and marine sediments from the Indian

(Kroon et al., 1991; Prell et al., 1992; Prell & Kutzbach,

1997) and North Pacific Oceans (Rea et al., 1998).

The summer conditions of the late Pliocene of the

Zhangcun region might be evaluated by the MWMT

and MMaP estimated by CA while winter conditions by

the MCMT and MMiP.

Our research showed that the summer temperatures

represented by the MWMT (Fig. 7b) remained constant

from Zone I to Zone IV while the summer precipitations

represented by MMaP (Fig. 7f) declined from Zone I to

Zone IV, implying a weakening of the summer monsoon

in the Zhangcun region during the late Pliocene. At the

same time, both winter temperatures represented by

the MCMT (Fig. 7c) and precipitations by the MMiP

(Fig. 7g) from Zone I to Zone IV declined, which might

have been triggered by a strengthening of the winter

monsoon.

The summer conditions of the late Pliocene Zhangcun

site were no doubt influenced by the southeasterly

summer monsoon from the Pacific Ocean since the

northward migration of southwesterly warm and wet

air from the Indian Ocean was probably blocked by the

height and extent of the Tibetan Plateau.

Table 3 The data of palaeoclimatic parameters of palynoassemblage as a whole and each of the four pollen zones using the

Coexistence Approach

Total Zone 1 Zone 2 Zone 3 Zone 4

MAT ( 1C) 8.5–15.1 8.5–20.9 8.5–15.1 7.8–22.1 7.8–22.7

MWMT ( 1C) 19.8–27.5 19.8–27.5 19.8–27.5 19.8–27.5 19.8–27.5

MCMT ( 1C) �0.3 to 2.0 �0.3 to 5.9 �0.3 to 2.0 �8.8 to 5.9 �8.8 to 5.9

DT ( 1C) 25.0–26.0 15.8–26.0 25.0–26.0 13.0–33.1 13.0–33.1

MAP (mm) 845.6–1050.9 845.6–1254.7 845.6–1050.9 573.9–1254.7 573.9–1254.7

MMaP (mm) 183.6–229.4 183.6–245.2 183.6–229.4 129.4–245.2 129.4–245.2

MMiP (mm) 19.2–21.2 19.2–21.2 19.2–21.2 5.7–21.2 5.7–21.2

Table 4 The comparison between the median values of seven

climatic parameters in the late Pliocene of Zhangcun and the

current meteorological data

Late Pliocene Modern*

MAT ( 1C) 11.8 8.8

MWMT ( 1C) 23.7 22.4

MCMT ( 1C) 0.9 �6.8

DT ( 1C) 25.5 29.2

MAP (mm) 948.3 578.9

MMaP (mm) 206.5 155.5

MMiP (mm) 20.2 4.6

*Refers to the record of Yushe Meteorological Station (371040N,

1121590E, 1041 m) about 15 km northeast of Zhangcun locality

(IDBMC, 1983a).



The winter monsoons in the late Pliocene Zhangcun

site would be strengthened by combining three kinds of

forces from (1) the northerly winter monsoon driven by

the Siberian High, (2) the northern of the two westerly

jet streams induced by the Tibetan Plateau, and (3) the

coalescence of the katabatic airflow caused by the

winter cold high pressure on the Tibetan Plateau with

the northerly airstream.

Zone(a)

I

II

III

IV

−8 −4 0 4 8

Zone

12

(c)

I

II

III

IV

18 20 22 24 26 28

(b)

I

II

III

IV

Zone

14 18 22 26 30 34

(d)

I

II

III

IV

Zone

130 170 210 250

(f)

I

II

III

IV

Zone

2 6 10 14 18 22MMiP (mm)

MAP (mm) MMaP (mm)

MCMT(°C) DT(°C)

MAT(°C) MWMT(°C)

Zone(g)

I

II

III

IV

600 800 1000 1200

(e)

I

II

III

IV

Zone

2 6 10 14 18 22

Fig. 7 Climatic parameter values of individual horizons (Zones I–IV) estimated by the Coexistence Approach. (a) The mean annual

temperature. (b) The mean warmest monthly temperature. (c) The mean coldest monthly temperature. (d) The temperature difference

between coldest and warmest months. (e) The mean annual precipitation. (f) The mean maximum monthly precipitation, (g) the mean

minimum monthly precipitation.



Acknowledgements

The authors thank Senior Engineer Nai-Qiu Du, Institute ofBotany, CAS, Beijing, China, for her kind help in identifyingthe pollen and spores, Dr. Ya-Meng Li, College of Life Science,Capital Normal University, Beijing, China, for valuable discus-sions, Dr. Nilamber Awasthi, Ex-Deputy Director, Birbal SahniInstitute of Palaeobotany, Lucknow, India, for reading the manu-script and useful suggestions during his visit to the Institute ofBotany, CAS, Beijing. This research was supported by the Na-tional Natural Science Foundation of China (Nos. 30770148;30990241; 30530050; 41072022), and Beijing Academy of Scienceand Technology (No. IG200704C2).

Author contributions: Yufei Wang and Chengsen Li conceivedthe ideas; Feng Qin and Yufei Wang collected the samples; FengQin and Svetlana Syabryaj identified the pollen & spores andanalysed the data; Reinhard Zetter analysed and identified the

pollen & spores and supplied the SEM plates; Feng Qin andYufei Wang wrote the first draft of this manuscript; Chengsen Li,Jinfeng Li, and Jian Yang revised the draft versions. David K.Ferguson rewrote some of the discussion and corrected the finalmanuscript.

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median of each climatic parameter.



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