动物学报 50 ( 5) :841 - 851 , 2004
A ct a Zoologica S i nica Relict distribution of land vertebrates and Quaternary glaciation in
China
ZHAN G Rong2Zu 3
Institute of Geographic Sciences and Natural Resources Research , CAS , Beijing 100101 , China
Abstract The biogeograp hical relict p henomenon of extant land vertebrates at t he species level has been considered in re2
lation to climatic changes since t he Quaternary. Three major relic patterns occurring in China have been illustrated. The
results seemly are not in agreement wit h t he t heory of t he East China Glaciation , a longstanding controversy [ A cta Zoo2
logica S i nica 50 ( 5) : 841 - 851 , 2004 ] .
Key words Biogeograp hical relict , Land vertebrates , East China Quaternary glaciation
中国第四纪冰期与陆生脊椎动物残留分布 张荣祖
中国科学院地理科学与资源研究所 , 北京 100101
摘 要 在第四纪冰期中 , 中国东部地区是否发生过冰川 , 长期存在争议 。中国东部地区冰川说的创始人李四
光教授在提出该学说时 , 从古生物方面未得到冰期生物群的证据 。现存生物种的地理残留现象可反映地质历史
时期的气候变化 。本文作者依据中国陆生脊椎动物的北方残留 、热带 - 亚热带残留 、温带湿润带残留等现象 ,
推断 : ( 1) 在冰期的冷期中喜暖动物南迁时 , 中国东部地区是一避难地 ; ( 2 ) 在中国东部地区未见有典型冰川
残留种的存在 ; ( 3) 一些湿润种的间断残留分布与中国北方黄土及干旱气候的发展有密切的关系 。此结论似不
支持中国东部冰川说 [ 动物学报 50 ( 5) : 841 - 851 , 2004 ] 。
关键词 生物地理残留 陆栖脊椎动物 中国东部第四纪冰川
1 Int roduction
In mainland Asia , major Pleistocene glaciations
came into existence in t he nort h , minor glaciations in
t he Cent ral High Asia , where t he huge Tibetan
( Qinghai2Xizang) Plateau lies and t he highest moun2
tain ranges of t he world , t he Himalaya , Karkorum ,
Kunlun , Tianshan in t he West China occur. Opinions
have varied greatly concerning t he extension of t he
glaciated areas in Siberia ( Nilsson , 1983 ) . Whet her
or not Quaternary glaciation ever existed in t he medi2
um and low mountains of East China is a long disput2
ed question. The famous Chinese geologist , Prof . Li
Si2Guang ( J . S. Lee ) and his followers claim t hat
glacial relics were found at over 100 places , even on
t he hilly land 200 m a1s1l. at 22°
N in East China. A
group leaded by Prof . Shi Yafeng came to different
conclusions. The main point is t hat apart f rom High
Asia , where t he glaciers have developed distinctly ,
Received Mar. 03 , 2004 ; accepted J uly 08 , 2004
3 ZHAN G Yong2zu
ν 2004 动物学报 A cta Zoologica S i nica
glaciers existed mostly in a few high mountains along
t he eastern margin areas of Tibetan Plateau. Moun2
tains lower t han 2 000 m over t he area of East China
were beyond t he condition to glaciers during Quater2
nary ( Fig11) ( Shi et al. , 1987) .
In an early paper on East China Quaternary
Glaciation , Prof . Li Si2 Guang ( 1942) recognized t hat
“based on palaontological evidence , ot her t han a mix2
t ure of cold , temperate or t ropical faunal element s , a
particular glacial biota has not been known in t he
glaciated area hypot hesized ” ( Li , 1975 ) . Then , a
t heory of Ice age ref ugia —
“ Pant u Ground ”of t he
East China was proposed ( Kahlke , 1961 ) . Recent
st udies on Quaternary palynology , mammals , loess ,
paleosols and cave deposit s show t hat t he climate os2
cillated f rom temperate to subt ropical many times
during t he Quaternary in Nort h China. A small num2
ber of relics of Mesozoic plant s , as ginkgo Ki nkgo
bitoba , water fir Met asequoia gly ptost roboi des and a
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动 物 学 报
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Fig11 Distribution of partial Quaternary glacier relics in Eastern China( Shi et al. ,1987)
large number of Tertiary plant s had been growing for
a long time and survived several Quaternary cooling
periods ( Shi et al. , 1987 ) . The Yangze crocodile
A lli gator si nensis and t he giant salamanders A n d ris
davi dian us , which are judged as species of t he t her2
mop hilic Tertiary fauna ( Illies , 1974 ) , are still ex2
tant in t he lower reaches of Changjiang River and
over t he most part of East China , respectively. The
record of t he Pleistocene zoogeograp hical evolution
and it s existing tendency indicate t hat t here was a
broad faunal t ransition zone between Oriental and
Palaearctic realms in East China ( Zheng and Han ,
1991 ; Zhang , 2002) . The result s of t hese st udies are
in essential agreement wit h t he t heory of ref ugium
rat her t han of glacial land in East China.
The synt hesis presented herein is based on inter2
pretations of relict dist ribution of extant land verte2
brates in China. It is an attempt to assemble impres2
sions accumulated during t he aut horπs long term st ud2
ies on extant land vertebrate dist ribution of China
t hat t he relic p henomenon could serve as evidence for
paleoenvironment changes ; and providing information
for t he of East China Glaciation cont roversy.
The concept of relict ness in t his paper is t hat of a
biogeograp hical relict ( Udvardy , 1969 ) , which oc2
curs in isolation f rom it s main center of dist ribution
induced by climatic change. Two examples , well
known in t he biogeograp hical literat ure , are t he
azure2winged magpie Cyanopica cyana and t he t hree2
toed woodpecker Picoi des t ri dactyl us , which have
been defined as temperate woodland relict and boreal2
alpine relict respectively , induced by t he vicissit udes
of t he Pleistocene climatic changes wiped out t he
species f rom t he connecting areas ( Voous , 1960 ,
quoted f rom Udvardy ,1969) . Their relic areas range
separately in China and t he Iberian Peninsula
( Fig12 ) . The following examples of most distinct
relic patterns , which are characterized by disjunctive
dist ribution at species level , would be obviously relat2
ing to t he question arose in terms of evolution time in
t he Quaternary. In order to focus t he question con2
cerning to climatic fluct uation , big game mammals
are not be included in t his paper. A list of 87 species
wit h t hose patterns has been attached ( see Ap2
pendix) .
The basic information of t he animal dist ribution
data of t his paper have been quoted f rom following
major cont ributions : ( 1 ) Amp hibians and reptiles by
Fei ( 1999 ) , Zhao and Adler ( 1993 ) , and Vorobev
(1988) ; ( 2 ) Birds by Cheng ( 1987 ) , Dementev et
al. ( 1951 - 1954 ) and Zheng ( 2002 ) ; ( 3 ) Mam2
mals by Zhang et al ( 1997 ) , Wang ( 2003 ) , Corbet
( 1978) , Sokolov and Orlov ( 1980) , Panteleev et al.
( 1990) and Flint et al. ( 1965) .
2 Boreal2alpine relict
During climatically induced faunal displace2
ment s , disjunctions f requently developed in t he cold2
adapting boreal2alpine species of Holarctica ( Illies ,
1974) . Similar dist ribution patterns are known f rom
5期
ZHAN G Rong2Zu : Relict distribution of land vertebrates and Quaternary glaciation
843
Fig12 Ranges of 1 , three2toed woodpecker Picoides t ridactyl us ; 2 ,azure2 winged magpie Cya nopica cya na ; 3 , Ussuri large
white2toothed shre w Crocidura lasi ura
China. Sout hwards shift of t he boreal fauna extended
as far as t he Cent ral China mountains is indicated by
number of species which have remained relic popula2
tion t here after t he nort hwards ret reat during t he
course of climatic fluct uation in t he Quaternary. Ex2
amples of t his include t he by following species :
( 1 ) Amp hibians Siberian salamander S al a2
m an d rell a keyseli n gii : it s main range coincides al2
most completely wit h t he area of taigas , and extent s a
bit f urt her to t he nort heastern part of t he Nort heast
China at about 45°
N . A relic population of t he species
has been recorded f urt her sout h , t he Tongbai2Dabie
mountains at about 32°
N in Henan province , Cent ral
China ( Fig13) ( Wu and Qu , 1984) .
( 2 ) Reptiles Adder V i pera berus : t he species
is dist ributed mainly in t he sout hwestern part of t he
taiga including nort hern part of Nort h East China and
Altai mountains , Xinjiang and f urt her to t he decidu2
ous forest zone of Cent ral Europe. A few records of
relic population has been known to t he sout h of Qin2
ling mountains at about 33°N in Cent ral China
( Fig13) ( Song , 1989) .
( 3) Birds Ural owl S t ri x u ralensis : t he main
range of t he species crosses f rom nort hern Europe
t hrough Siberia reaching t he coast of t he Far East and
including Sakhalin Island and J apanese islands. A rel2
ic population occurs in t he nort hern Hengduan moun2
tains at 30 - 34°N in Sout hwest China ( Fig14 )
( Cheng , 1987) .
( 4) Mammals Root vole M icrot us eoconom us :
t he main range of t he species covers most of t he taiga
and much of t he t undra in t he nort h and extend to t he
Cent ral Mongolia and Tianshan mountains in t he
sout h where it spreads as alpine taiga and wooded
steppe. A relic population exist s in t he nort heastern
margin of t he Tibetan Plateau about 32 - 40°N
( Fig14) . The narrow2 skulled vole M icrot us gregalis
is dist ributed continually in t he taiga of Siberia and
spread sout hwards along t he mountain chains of Da
Xinganling and Tianshan in China. The relic popula2
tion of t he species occurs in Funiu mountains , about
32 - 35°
N in Henan Province.
3 Tropical2subt ropical relict
Palaeontological st udies show t hat prior to t he
first cold period of t he Quaternary , a t hermop hilic
fauna most belonging to groups now occurring in t he
Oriental region associated wit h t ropical2subt ropical
forest s , was dist ributed over most of China ( Zhou ,
1984) . As climatic deterioration progressed since t he
Pleistocene in China , t he species were forced far
sout hwards especially in t he cold stages ( Zheng and
Han ,1991 ; Ji , 1985 ) . In t he course , relict popula2
tions may have remained in t he nort h where ref uge
habitat s existed , while t he main ranges ret reated to
t he areas reaching lower latit udes in different dis2
tance. The following species may be examples :
( 1 ) Amp hibians Large odorous f rog
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动 物 学 报
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Fig13 Ranges of 1 , Siberian salamander Salama ndrella keyselingii ;2 ,adder Vipera berus ; and 3 , boreal dig2
ging frog Kaloula borealis
Fig14 Ranges of 1 , Ural owl St rix uralensis ; and 2 ,root vole Microt us eoconomus
( O dorrana li vi da = R ana li vi da ) maintains it s relic
population in t he sout heastern Shaanxi , to t he sout h
of Qinling at about 34°
N , while t he nort hern limit of
it s main range is laying to t he sout h of t he Changjiang
River ( Fig15) ( Fei ,1999) .
( 2 ) Reptiles Green pit2viper T ri meresu rus
stej negeri has a relic population recorded in Changbai
Mountain at about 42°
N in Nort h East China , where
5期
ZHAN G Rong2Zu : Relict distribution of land vertebrates and Quaternary glaciation
845
Fig15 Ranges of 1 ,large odorous frog Ra na livida ; 2 ,green pit2viper Trimeresurus stej negeri ; and
3 , rhesus monkey Macaca mulatta
a temperate zone is sit uated ( Zhao , 1979) ( Fig15) .
These Oriental species are mainly dist ributed in
sout heast Asia , wit h a nort hern limit approximately
coincided wit h t he nort hern boundary of t he nort h
subt ropical zone. In cont rast to t he green pit2viper ,
t he Banded coral snake Calliophis m acclell an di has a
relic population in t he border area at about 33°N in
Gansu and Shaanxi ( Zhao and Adler , 1993 )
( Fig16) , along t he most nort hern limit of nort h sub2
t ropical zone and , t he nort hern limit of it s main range
is bounded by t he cent ral subt ropical zone.
( 3) Mammals Sout h China hare L epus si nensis
dist ributed mainly in cent ral subt ropical zone of East
China , a relic population record has been known at
about 42°
N in Changbai mountain , Nort h East China
( Fig16) (L uo , 1988) . Probably it belongs to t he rel2
ic population of t he same species in t he Korean penin2
sula. An isolated population of rhesus monkey M aca2
ca m ul att a occurred at about 41°N in t he nort h of
Beijing , which population was at t he nort hernmost
edge of t he species in Asia. A few individuals of t he
population disappeared due to human impact in t he
end of 1980s ( Fig15 ) ( Zhang et al. , 1989 ) . Now ,
t he main range of t his species in t he area of subt ropi2
cal zone in China is more or less continuous in terms
of geograp hical dist ribution on a small scale map , an
insolated population laying just nort h of t he Yellow
River at about 35°N in border area of Henan and
Shanxi. The black giant squirrel R at uf a bicolor
could be an example of t hose species , which have re2
t reated fart her sout h. The main range of t his species
is in t he Indo2China Peninsula , but remains on a mar2
gin of it in sout hern Yunnan Province and a relic pop2
ulation in Hainan island ( Fig16 ) bot h belong to t he
t ropical zone of China.
4 Temperate2humid relict
Deposition of loess indicating a tendency of
change toward a of dry and cold climate , began in t he
early Pleistocene , but it s dist ribution was limited to
t he nort h of t he Qinling mountains in China. By t he
Late Pleistocene , it had become much more
widespread , reaching as far as t he middle and lower
reaches of t he Changjiang River. The widespread de2
position of loess dist urbed t he previous humid forest
land ( Liu and Ding , 1984 ) . The species of st rict
mesop hytes , adapted to humid2temperate habitat s
disappeared in t he main part of t he deteriorated areas
and may have remained relic population scattered in
locally suitable habitat s out side or inside of t he arid
land. Examples of t his pattern are given by following
species :
(1 ) Amp hibian Boreal digging f rog Kaloul a
borealis is dist ributed mainly around t he eastern part
of nort h China , a semi2humid area , while a relict
population occurs in Wenxian of sout hwestern Gansu
Province ( Fig13) ( Fei et al. , 1999) , where is wit h2
in t he margin of t he humid subt ropical zone.
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动 物 学 报
50 卷
Fig16 Range of 1 , banded coral snake Calliophis macclella ndi ; 2 , South China hare Lepus sinensis ; 3 , black giant
squirrel Rat uf a bicolor
( 2) Reptiles Taky d rom us am u rensis is mainly
dist ributed in t he eastern part of Nort h East China ,
but a relic population has been recorded in sout heast2
ern Gansu Province ( Fig17 ) ( Yao and Chang ,
1981) , in a location near t he relict population of bo2
real digging f rog ( see above) .
(3 ) Birds t he Reed parrot bill Paradoxornis
heu dei has two separate ranges , one of t hem in t he
nort h , t he cent ral area of Nort h East China , t he an2
ot her in t he sout h , in Zejiang Province ( Fig17 )
( Cheng ,1987) .
( 4 ) Mammals J apanese mole M ogera w ogu ra
is mainly dist ributed in t he Korean Peninsula ,
Liaodong peninsula and J apanese islands. Two relic
populations occur in lower reaches of Yellow River
and Changjiang River ( Fig17 ) . Chinese birch mouse
S icist a concolor has five separate relic ranges located
in mountain areas : Sikhote Alin2Sungari in t he east ,
Nanshan2Qinling , West Tienshan , Hindu Kush and
Caucasus in t he west ( Fig17 ) ( Flint et al. , 1965 ;
Corbet , 1978 ; Zhang et al. , 1997 ) . Ussuri large
white2toot hed shrew Croci d u ra l asi u ra has a main
range in Korea Peninsula2Ussuri area and eastern
mountains of Nort h East China , and one relic popula2
tion occurs in lowest reaches of Changjiang River ,
( Fig12 ) separated by vast arid2semihunid land. De
Wintonπs shrew S oricul us hy psibi us is mainly dis2
t ributed in t he mountains st retching along t he nort h2
ern margin of Sichuan Basin and a relic population oc2
curs near Beijing ( Fig17) .
5 Discussion
During t he last glaciation of t he Quaternary ,
stimulated by glacial episodes , arctic t undra faunas e2
volved in t he nort hern hemisp here. A few mammals
of st rongly t undra2specific , t he polar bear U rsus
m ariti m us , polar fox A lopex l agopus , reindeer
R angif er t aran d us , collared lemming Dicrostony x
and narrow2skulled vole M icrot us gregalis are repre2
sented. To t he sout hern f ront of t he glacial land , t he
region of coniferous forest , t he taiga , which t hough
not quite as ext reme climatically is still characterized
by a distinctly low temperat ure. The fauna of t he
taiga includes all t he t undra animals , but wit h t he ad2
dition of most of more temperate latit udes ( Illies ,
1974 ; Marain , 1984 ) . Among t hem t he wolverine
Gulo gulo and polar hare L epus ti m i d us st retch a lit2
tle sout hwards , reaching t he nort hernmost area of
China , associating wit h t he taiga zone in Nort h East
China and Altai mountains , Xinjiang. The root vole
M icrot us oeconom us and narrow2 skulled vole M icro2
t us gregalis spread much f urt her to t he sout h and re2
mained relic populations. The relic range of root vole
occurs in t he nort heastern margin of Tibetan Plateau
where periglacial habitat s and alpine glaciers still ex2
ist . The relic range of narrow2skulled vole occurs in
much lower elevation where is wit hin t he area of
Pleistocene glaciation suggested by t he scholar Prof .
5期
ZHAN G Rong2Zu : Relict distribution of land vertebrates and Quaternary glaciation
847
Fig17 Ranges of 1 , Takydromus amurensis ; 2 , De Wintonπs shre w Soricul us hypsibi us ; 3 , Chinese birch mouse
Sicista concolor ; 4 ,Japanese mole Mogera wogura ; and 5 ,reed parrot bill Pa radoxornis heudei
Li. So far known , in Cent ral Europe , t hese two
species of voles survived t he individual stadials at t he
lower elevations , denoted warmer periods , character2
ized by a disjunct occurrence , differing f rom typically
nort hern t undra species ( Povolny , 1966) . No doubt ,
t he relics of t he two species of voles and ot her taiga
faunal species would be t he evidence for t he experi2
ence of relatively cold climate of t he areas in t he past
when t he taiga extended sout hwards , but not neces2
sarily under conditions of glaciation. And , even much
of Siberia , where t he taiga originated , shows no geo2
logic evidence of glaciation in t he Pleistocene (Brown
and Gibson , 1983 ) . Obviously , t he changes of
sout hern limit s ranging f rom t he boreal relic in China
may indicate t he extent of cold climate during cold
stages of t he Pleistocene. The animals of t he taiga
could migrate sout hwards at least along cold ridges
wit h alpine coniferous of t he mountains in nort hern
China. The occurrence of t he boreal - alpine relic
species have reached mainly to t he Qinling2Dabie
mountains in t he east and to eastern margin of t he Ti2
betan Plateau , t he Hengduan mountains , in t he
west , between 30 - 40°
N.
During maximum glaciation , t he Tibetan Plateau
was never covered by ice sheet s , but under periglacial
environment wit h different scale of valley2pediment
glaciers mainly ( Shi and Zheng , 1995) . Palynological
st udies show t hat since t he Pliocene t he general t rend
of environmental changes of t he plateau has been f rom
warm humid to cold arid climate and t he forest s have
ret reated f rom t he interior to t he sout heastern margin
in lower elevation. In t he course of t he uplift of t he
plateau , alpine t undra faunas have evolved and
formed a distinct highland pattern in dist ribution
( Hsü, 1978 ; Zhang and Zheng , 1981 ) . Among
mammals , t he black2lipped pike Ochotona cu rzoniae ,
Himalayan marmot M arm ot a hi m al ayana , Blyt hπs
vole Pity m ys leucu rus white 2lipped deer Cerv us al 2
bi rost ris , etc. are specialized for periglacial alpine
meadow habitat s. Their fossils have been known
firstly in sout hern Tibet in t he st rata of early
Holocene ( Guo , 1976) . Neit her fossil nor relic popu2
lation of t hem have been known f rom East China , but
t heir ranges extend to t he eastern margin of t he Ti2
betan plateau , around Hengduan mountains , where
glaciation has been confirmed occurring in few high
mountains during t he Pleistocene , even now.
Before t he middle Pleistocene , t he subt ropical
zone extended nort hwards , reaching to t he sout h at
42°
N in Nort h East China ( Liu , 1984 ) . Associated
wit h t he climatic optimum , t he nort hern dist ribution
limit of t he warm2preferring animal species during
t hat time could be shown by t he nort hernmost loca2
tions of t heir relics. A few of t hem occur at latit ude of
42°
N to t he nort h of t he Korea Peninsula , coinciding
wit h t he nort hern limit of middle Pleistocene subt rop2
ical zone and is close t he present nort hern margin of
warm2temperate zone. During sout hwards ret reat s of
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动 物 学 报
t he subt ropical zone in t he cold stage since middle
Pleistocene , numbers of t he warm2preferring species
remained relic population in different places depend2
ing on t heir ability to adapt to t he cold2resistant and
habitat conditions in t he areas ret reated. The nort h2
ern limit s of t he main ranges of t hose species lie at
different latit udes wit h various contours coinciding
wit h suitable optimal environment s. Some of t hem
have wit hdrawn f rom t he subt ropical areas or only re2
main in t he islands of Hainan and Taiwan , even out
of t he territory of China. The t ropical2subt ropical
relics could indicate t hat in East China since t he Mid2
dle Pleistocene to t he Later Pleistocene , t he climatic
conditions have fluct uated f rom temperate to subt rop2
ical wit hin t he limit of cold2tolerant of t he warm2pre2
ferring relic species.
The progressive develop ment of arid climate and
loess deposition as mentioned above resulted in an arid
land of so called“Nort h West Arid Region”in China ,
which includes desert and steppe. To t he east , t he
region connect s t he forest2steppe region of t he Nort h
China and forms a relatively arid zone crossing nort h2
ern China. The former scale of t he zone could be indi2
cated by dist ribution of a few representative species ,
such as sand lizards ( Phry nocephal us spp . ) in rep2
tiles and zokors ( M yos pal ax spp . ) in mammals ,
which range continuously over most land of Nort h
China , eastern Inner Mongolia and t he east of Nort h
East China ; no data in terms of biogeograp hical relic
have been learned f rom t hem. On t he cont rary , t he
remarkable p henomenon of t he extant widely discon2
tinuous gaps of t he humid2temperate relic species
( Fig17) could not be originated by crossing t he barri2
er of Po Hai Sea or vast loess plateau , except t he pos2
sible migration by chance of t he reed parrot bill , but
because t he population disappeared in t he Nort h Chi2
na plain , and point s at a general tendency of intensive
climatic deterioration of cold and arid wit h minor
fluct uation since early Pleistocene during t he evolution
of t he loess plateau. The same could be t rue of all t he
ot her two cases of mesop hytic species. Obviously ,
arid condition is one of t he most important negative
factors for glacier formation.
6 Conclusions
This elucidation suggest s t hat since t he Quater2
nary , t he following event s could be identified. ( 1 )
Eastern China has become a ref ugium for survival of
temperate and t ropical2subt ropical faunas during t heir
sout hwards ret reat in t he cold stage ; ( 2) a number of
taiga fauna have extended to t he middle reaches of
Changjiang River , but no typically glacial relic has
been found in t he lower lands of East China ; ( 3 ) a2
mong t he faunal species of sout hwards2 nort hwards
extension associated wit h taiga and temperate forest s ,
50 卷
t he numbers of mesop hytic specialized species have
been become discontinuous in dist ribution by t he de2
velop ment of t he loess arid and semi2arid zone. This
conclusion seemly is not in agreement wit h Prof . Liπs
hypot hesis of East China Glaciation.
Acknowledgements I would like to express my deep2
est appreciation to Prof . Shi Ya2Feng , Prof . Zeng
Zhao2Xuan and t hree anonymous referees for t heir
comment s.
References
Brown J H , Gibson A G , 1983. Biogeography. St . Louis : The C1V.
Mosby Co.
Cheng TH , 1987. A synopsis of t he Avifauna of China. Beijing : Sci2
ence Press , Hamburg and Berlin : Paul Parey Scientific Publishers.
Corbet GB , 1978. The Mammals of t he Palaearctic Region : A Taxo2
nomic Review. London : Cornell University Press.
Dementiev GP , Gladkov NA , 1951 - 1954. Birds of Soviet Union Vol.
1 - 6. Moscow : Science Press ( In Russian) .
Fei L , 1999. Atlas of Amphibian of China. Zhengzhou : Henan Science
and technical Press ( In Chinese) .
Flint V E , Chygynov UD , Smirin VM , 1965. Mammals of USSR.
Moscow : Meisle Press ( In Russian) .
Guo XD , 1976. Quaternary interglacial period and Paleoclimate in t he
Area of Zhumulangma Pike. In : Scientific Expedition Team , Chi2
nese Academy of Sciences ed. The Scientific Expedition Reports of
t he Area of Zhumulangma Pike (1966 - 1968) : Quaternary Geolo2
gy. Beijing : Science Press : 63 - 78 ( In Chinese) .
HsüJ , 1978. On t he palaobotanical evidence for continental drift and
Himalayan uplift . Palaeobotanist 25 : 131 - 142.
Illies J , 1974. Introduction to Zoogeography. London : The Macmillan
Press Ltd , 57.
Ji HX , 1985. Mammal distribution and its reflection on climatic changes
in t he Quaternary of China. In : Proceedings of Symposium on
Glacial and Periglacial China ( 1985 ) . Beijing : Science Press ,
138 - 142 ( In Chinese) .
Kahlke HD , 1961. On t he complex of t he S tegrodon2A il uropoda fauna
of Sout hern China and t he chronological position of Giant pit hecus
blacki V. Koenigswald. Vertebrata PalAsiatica 2 : 85 - 103 ( In
Chinese) .
Li SG , 1975. Inquiry into Glaciation of China. In : Li SG ed. Quater2
nary Glaciation of China. Beijing : Scientific Press , 118 - 124 ( In
Chinese) .
Liu DS , Ding ML , 1984. The characteristics and evolution of t he
Palaeo environment of China since t he Late Tertiary. In : Whyte
RO ed. The Evolution of t he East Asian Environment . Vol. Ⅰ:
Geology and Palaeoclimatology. Hong Kong : University of Hong
Kong , 12 - 40.
L uo ZX , 1988. The Chinese Hare. Beijing : China Forestry Publishing
House , 85 ( In Chinese) .
Marain A , 1984. Systematic and Regional Biogeography. New York :
Van Nostrand Eeinhold Company , 212.
Nilsson T , 1983. The Pleistocene —Geology and Life in t he Quaternary
Ice Age. Dordrecht , Boston , London : D. Reidel Publishing Co. ,
311.
Panteleev PA , Terexina AN , Varshavskii AA , 1990. Ecogeographical
Change of Rodents. Moscow : Science Press ( In Russian) .
Povolny D , 1966. The Fauna of Central Europe : Its Origin and Evolu2
tion. Systematic Zoology 45 : 46 - 53.
Shi YF , Zheng BX , 1995. Timing and height of t he Qinghai2Xizang
Plateau uplifting into t he cryosphere and its impact on t he sur2
rounding areas. In : Formation , Evolution , Environment Changes
and Ecosystem of Qinghai2Xizang Plateau. Beijing : Science Press ,
136 - 146 ( In Chinese) .
Shi YF , Cui ZJ , Li JJ , 1987. Reassessment of Quaternary glaciation
5期
ZHAN G Rong2Zu : Relict distribution of land vertebrates and Quaternary glaciation
problems in East China. Eart h Sciences 2 : 45 - 54.
Sokolov V E , Orlov VN , 1980. Mammal Determination of Peopleπs Re2
public of Mongolia. Moscow : Science Press ( In Russian) .
Song M T , 1989. The Herpetofauna of Shaanxi Province. Acta Her2
prtologica Sinica 6 (4) : 63 - 73 ( In Chinese) .
Udvardy MDF , 1969. Dynamic Zoogeography. New York , Van Nos2
trand : Reinhokd Co . , 205.
Vorobeav ES , 1988. Amphibians and Reptiles of Mongolian Peopleπs
Republic. Moscow : Science Press ( In Russian) .
Wang YX , 2003. A Complete Checklist of Mammal Species and Sub2
species in China. Beijing : Forestry Publishing House ( In Chi2
nese) .
Wu SH , Qu W Y , 1984. A Preliminary study of amphibian fauna of
Henan Province. Journal of Xinxiang Normal College 41 : 89 - 93
( In Chinese) .
Yao CY , Chang SZ , 1981. Three new records of reptiles from Gansu.
Acta Herpetologica Sinica 5 (9) : 65 - 68 ( In Chinese) .
Zhang YZ , Jin SK , Li SH , Ye ZY , Wang F G , ZHang ML , 1997.
Distribution of Mammalian Species in China. Beijing : China
Forestry Publishing House.
Zhang YZ , Quan GQ , Lin YL , Sout hwick C , 1989. Extinction of rhe2
sus monkeys M acaca m ulatta in Xinglung , Nort h China. Interna2
tional Journal of Primatology 10 (4) : 375 - 381.
Zhang RZ ( YZ) , 1999. Zoogeography of China. Beijing : Science
Press ( In Chinese) .
Zhang YZ , Zheng CL , 1981. The geographical distribution of mammals
and t he evolution of mammalian in Qinghai2Xizang Plateau. In :
Geological and Ecological Studies of Qinghai2Xizang Plateau. Vol.
Ⅱ: Environment and Ecology of Qinghai2Xizang Plateau. Beijing :
Science Press : 1 005 - 1 002.
Zhang RZ ( YZ) , 2002. Geological events and mammalian distribution
in China. Acta Zool. Sin. 48 (2) : 141 - 153.
Zhao EM , Adler K , 1993. Herpetology of China. Oxford , Ohio :
Society for t he Study of Amphibians and Reptiles.
Zhao EM , Yan ZK , 1979. Newly found green pit2viper in Changbai
Shan , Jilin , and discussion on its geographical distribution. Acta
Herpetologica Sinica 1 : 1 - 3 ( In Chinese) .
Zheng GM , 2002. A checklist on t he Classification and Distributuion of
t he Birds of t he World. Beijing : Science Press.
Zheng SH , Han DF , 1991. Quaternary mammals of China. In : Liu
TS ed. Quaternary Geology and Environment in China. Beijing :
849
Science Press , 101 - 114.
Zhou TR , 1984. Physico2geography2Palaeogeography ( Part Ⅰ) . Bei2
jing : Science Press , 74 - 133 ( In Chinese) .
费 梁 , 1999. 中国两栖动物图鉴 . 郑州 : 河南科技出版社 .
郭旭东 , 1976. 珠穆朗玛峰地区第四纪间冰期和古气候 . 见 : 中国
科学院西藏科学考察 队 编 . 珠 穆 朗 玛 峰 地 区 科 学 考 察 报 告
(1966 - 1968) : 第四纪地质 . 北京 : 科学出版社 : 63 - 78.
计宏祥 , 1985. 中国第四纪哺乳动物群的地理分布及其所反映的气
候变迁 . 见 : 中国第四纪冰川冰缘学术讨论会文集 . 北京 : 科
学出版社 : 138 - 142.
Kahlke HD , 1961. 关于中国南方剑齿象 —
—
—熊猫动物群和巨猿的
时代 . 古脊椎动物与古人类学报 . 2 : 85 - 103.
李四光 , 1975. 中国冰期之探讨 . 见 : 李四光著 . 中国第四纪冰
川 . 北京 : 科学出版社 : 118 - 124.
罗泽　, 1988. 中国野兔 . 北京 : 中国林业出版社 , 85.
施雅风 , 郑本兴 , 1995. 青藏高原进入冰冻圈的时代 、高度及其对
周围地区的影响 . 见 : 青藏高原形成 、演化 、环境变迁与生态
系统 . 北京 : 科学出版社 : 136 - 146.
宋鸣涛 , 1989. 陕西两栖爬行动物区系分析 . 两栖爬行动物学报 6
(4) : 63 - 73.
王应祥 , 2003. 中国哺乳动物种和亚种分类名录与分布大全 . 北
京 : 中国林业出版社 .
吴淑辉 , 瞿文元 , 1984. 河南省两栖动物区系初步研究 . 新乡师范
学院学报 41 : 89 - 93.
姚崇勇 , 张绳祖 , 1981. 甘肃爬行类三种新记录 . 两栖爬行动物研
究 5 (9) : 65 - 68.
张荣祖 , 1999. 中国动物地理 . 北京 : 科学出版社 .
赵尔宓 , 严仲凯 , 1979. 竹叶青蛇在长白山的发现及其地理分布的
探讨 . 两栖爬行动物研究 1 : 1 - 3.
周廷儒 , 1984. 中国自然地理 —
—
—古地理 ( 上册) . 北京 : 科学出版
社.
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50 卷
Appendix : Biogeographical relic list of extant land vertebrates in China
1. Boreal2alpine rel ict
A : Siberian salamamder S al am am d rell a keyserli n gii ( To ) .
R : adder V i pera berus ( Qs) .
B : Chinese grouse B onasia bonasia ( H) , boreal owl A egoli us f u nereus ( T) , long2eared owl Asio ot us ( T) , U2
ral owl S t ri x u ralensis ( T) , t hree2toed woodpecker Picoi des t ri dactyl us ( H) , white2backed woodpecker
Picoi des leucotos ( H , Fuj , It ) , Siberian rubyt hroat L usci nia calliope ( H ) , dusky warbler Phylloscopus
f uscat us ( H ) , Pallasπs leaf warler Phylloscopus p roregul us ( H ) , yellow2browed warler Phylloscopus
p roregul us ( H ) , red crossbill L ox ia cu rvi rost ra ( H ) , common rosefinch Carpodacus eryt hri n us ( H ) ,
Chinese nut hatch S itt a vilosa ( T , wt ) .
M : nort hern hedgehog Erianceus eu ropaeus ( subt . , Eu ) , east Siberian shrew S orex m i n utissi m us ( H ) ,
pygmy shrew S orex m i n ut us ( H) , Laxmannπs shrew S orex caecutiens ( T) , common shrew S orex araneus
( H , subt . ) , nort hern pika Ochotona al pi na ( wt ) , Siberian flying squirrel Pterom ys vol ans ( To ) , Eurasian
red squirrel Sci u rus v ul garis ( To ) , nort hern red2backed vole Clet heionom ys rutil us ( wt ) , narrow2skulled
vole M icrot us gregalis ( To ) , root vole M icrot us oeconom us ( T) , common vole M icrot us arv alis ( T) .
2. Tropical2subtropical rel ict ( most in t he nort hern margin of t he subt ropical zone)
A : black knobby newt Tylotot riton asperri m us ( To ) , bonyheaded toad B uf o galeat us ( Ih) , Sout h China t ree
toad Hyl a si m plex ( Z) , large odorous f rog ( Q ) red2webbed t reef rog R hacophorus rhodopus ( D) , beautif ul
pygmy f rog M icrohyl a pulchra ( W) .
R : T rionys stei n dachneri ( N . subt . ) , S acalia bealei ( C. , subt . ) , banded coral snake Calliophis m accell an di
( W) , A chali n us ruf escens ( Qs) , T rachischi u m ten uiceps ( Qs) , red2necked keelback R habdophis sub2
m i niat a ( wt ) , green pit viper T ri meresu rus stej negeri ( t ) , Pareas m onticol a ( Fj. subt . )
B : black2browed barbet Megal ai m a oorti ( D. , Ih2t ) , blue2rumped pitta Pitt a soror ( D , Ih ) , maroon oriole
O riol us t raillii ( Ih , It ) , black drongo Dicru rus m acrocercus ( H , Ih , It ) , bronzed drongo Dicru rus
aeneus ( H , Ih , It ) , greater racket2tailed drongo Dicru rus paradiseus ( Ih) , short2tailed green magpie Cis2
sa t hal assi na ( D , subt . , Ih) , spot2necked babbler S t achy ris st riol at a ( Ih) , yellow2browed tit S yl vi parus
m odest us ( H , Fj , Gus) , sultan tit Mel anochlora sult anea ( H , Gus , Fj , Ih) .
M : common European white2toot hed shrew Croci d u ra russul a , Hodgsonπs brown2toot hed shrew S oricul us cau 2
dat us ( H , Qs , It ) , Horsfieldπs shrew Croci d u ra horsf iel di ( Qs , H , Ih. t ) , Indian flying fox Pteropus
gi ganteus ( Qs. T) , greater false vampire Megaderm a ly ra ( n. subt ) , little J apanese horseshoe bat R hi2
nolophus corn ut us ( wt ) , Schreiberπs long2fingered bat M i niopterus schreibersi ( wt ) , wrinkle2lipped bat
Tadari da plicatt a ( wt ) , hairy2winged bat Harpiocephal us harpia ( H. It ) , Chinese hare L epus si nensis
( t . ) , rhesus macaque M acaca m ul atto ( wt , h) , black gibbon Hylobates concolor ( Ih) , hairy2footed flying
squirrel Belom ys pearsoni ( To ) , red giant flying squirrel Pet au rist a pet au rist a ( W , Qs , It ) , red2and2
white flying squirrel Pet au rist a alboruf us ( H , W , Qs , It ) , Himalayan st riped squirrel Tam iops m acclel 2
l an di ( To ) , black giant squirrel R at uf a bicolor ( Ih) , Chinese bamboo rat R hizom ys si nensis ( Qn) Chinese
porcupine Hyst ri x hodgsoni ( wt ) , palm mouse V an deleu ria oleracea ( H) , pencil2 tailed t ree mouse Chi 2
ropodom ys gli roi des ( D , Ih) , black rat R att us ratt us ( wt ) .
3. Temperate2humid rel ict ( relic population occurring in monsoon regions separated by loess2arid area. )
A : oriental bell taod bom bi na orient alis , boreal digging f rog Kaloul a borealis ( W) .
R : Taky d rom us am u rensis ( t . W) , Taky d rom us w olteri ( subt . , t . ) Taky d rom us septent rionalis ( Subt . ,
t. ) .
B : reed parrot bill Paradoxornis heu dei ( subt . ,t . ) .
M : Nattererπs bat M yotis natterei ( t . , ct . , Eu. ) , water bat M yotis daubentoni ( t . ,ct . , Eu. ) , long2fingered
bat M yotis capacci nii ( subt . , t ,. Eu. ) , particolored bat V es pertilio m u ri n us ( t . , ct . , Eu. ) , Chinese
water deer Hy d ropotes i nerm is ( subt . , t . ) Chinese birch mice S icist a concolor ( H , Tianshan , t . ) , Ussuri
large white2toot hed shrew Croci d u ra l asi u ra , De Wintonπs shrew S oricul us hysibi us .
A : amp hibian. R : reptile. B : birds. M : mammals.
Relic areas. ct : Cold temperate zone. D : Dayaoshan , Guangsi. Eu. Europe. Fj : Fujian. Gus : Guangxi. H :
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851
Hengduan mountains. Ih : Hainan Is. It : Taiwan Is. Q : Qinling mountains ( Qs : sout hern flank ; Qn : nort h2
ern flank) . subt . : Subt ropical zone. t : Temperate zone. T : Tibetan plateau. To : Tongbai2Dabie mountains ,
Henan. t r : Tropic zone. W : Wenxian , Gansu ( to t he west of Qinling mountains) . wt : Warm temperate zone.
Z : Zheijiang.