全 文 ：A multivariate morphometric study on Corylus sieboldiana
　 complex(Betulaceae)in China , Korea , and Japan 　
1CHANG Chin_Sung　1CHANG Gae_Sun　2QIN Hai_Ning
1(The Arboretum and Department of Forest Resources , Seoul National University , Suwon 441_744, Republic of Korea)
2(Institute of Botany , the Chinese Academy of Sciences , Beijing 100093 , China)
Abstract　Previous taxonomic treatments in Russia and China have considered Corylus mandshurica
and C.sieboldiana to be distinct as independent species.A morphometric analysis was conducted
to determine if the morphological differentiation from these taxa warrants specific taxonomic recogni-
tion with a large sample of field_collected leaves as well as specimens from several herbaria of north-
eastern Asia.One hundred and fifty two individuals representing Chinese , Korean , and Japanese
geographic ranges of the species were scored for 18morphological characters and the data matrix was
used for principal components analysis.The entities that comprise C.sieboldiana complex exhibit
widely overlapping ranges in all morphological attributes.The leaf and fruit data may reflect a lack
of divergence between taxa.Therefore , we do not regard these to be sufficient for taxonomic splitting
of C.mandshurica from C.sieboldiana to warrant the designation of the rank of a species based on
morphology.The two taxa were not morphologically well differentiated and their ranges of distribution
come together.Therefore , C.mandshurica should be recognized as an infraspecific taxon of C.
sieboldiana.C.sieboldiana with short involucres in southern Korea and Japan is often treated as an
independent species (C.hallaisanensis)or a variety (C.sieboldiana var.brevirostris), but
should be only recognized as a form of C.sieboldiana because this involucral character is highly
variable even within the same individual.
Key words　Corylus sieboldiana complex , morphometric principal components analysis , taxonomic
treatment.
Corylus L.is a wide_ranging genus of trees and shrubs comprising up to 15(Furlow , 1990)or
20(Rehder , 1927)species throughout the NorthernHemisphere with most of the species distributed
in Eurasia.Although many significant contributions on hazelnut phylogeny have been made in DNA
studies(Forest &Bruneau , 2000;Erdogan &Mehlenbacher , 2000;Whitcher &Wen , 2001),
most results have proposed only the broad outlines of a phylogeny for extant taxa.
Infrageneric classifications of Corylus have recognized either two or three main divisions as sec-
tions or subgenera (Furlow , 1990;Whitcher &Wen , 2001).The ITS phylogeny (Whitcher &
Wen , 2001)supported the recognition of two sections(Acanthochlamys Spach and Corylus)with
section Corylus being divided into three subsections (Corylus , Colurnae Schneid. and
Siphonochlamys(Bobrov)P.C.Li)(Forest&Bruneau , 2000).However , many taxonomists often
recognized two subsections within sect.Corylus primarily based on the morphology of the involucre
surrounding nut(de Candolle , 1864;Schneider , 1916;Furlow , 1997).Corylus sieboldiana Blume
and C.mandshurica Maxim.&Rupr.belong to sect.Corylus , subsect.Siphonochlamys(sensu
Furlow , 1997;Li &Cheng , 1979)due to tubular involucre.
　Received:27 May 2003　Accepted:5 March 2004.
Supported partly by a grant from Plant Diversity Research Center of 21st Frontier Research Program funded by Minist ry of Science
and Technology of Korean Government(Grant No.PF001302_00).
植 物 分 类 学 报　42(3):222-235(2004)
Acta Phytotaxonomica Sinica
Species delimitation in the C.sieboldiana complex has been discussed by several authors
(Schneider , 1916;Kitamura &Murata , 1984;Ohwi , 1984;Fu et al., 2000;Charkevicz ,
1996), but is still debatable.C.sieboldiana var.sieboldiana , C.sieboldiana var.brevirostris
Schneid., C.mandshurica , and C.hallaisanensis Nakai are here collectively referred to as the
C.sieboldiana complex.
After Blume first recognized C.sieboldiana in Japan , several infraspecific taxa , var.mitis
Nakai , var.brevirostris Schneid., and var.mandshurica (Maxim.&Rupr.)Schneid., were distin-
guished by the size of fruit , leaf shape , leaf length , and shape of involucre (Schneider , 1916;Kita-
mura &Murata , 1984;Ohwi , 1984).Maximowicz and Ruprecht(1856)considered C.mandshurica
as a distinctive species , while Schneider(1916)treated C.mandshurica at the infraspecific rank of
C.sieboldiana , i.e., C.sieboldiana var.mandshurica.Recent floras in Russia (Charkevicz ,
1996)and China(Li &Cheng , 1979;Li &Skvortsov , 1999;Fu et al., 2000)presented arguments
favoring recognition of this taxon as a species.However , Japanese and Korean taxonomists(Kitamura
&Murata , 1984;Lee , 1980;Ohwi , 1984;Lee , 1996)differed from Chinese and Russian botanists
in reducing C.mandshurica to a variety of C.sieboldiana.Two major treatments regarding C.
sieboldiana and C.mandshurica differed from each other primarily in their recognition of four mor-
phological characters:leaf shape , leaf serration , involucre width , and involucre length.C.
sieboldiana is characterized by its obovate_oblong leaves that are rounded at base and more or less
gradually pointed at apex with acute serration.On the other hand , C.mandshurica is characterized
by its orbicular_ovate leaves that are more truncate at apex and mostly coarser , shorter and broader
lobed above the middle.The involucre of C.sieboldiana is contracted above the nut into a narrower
tube , while that of C.mandshurica is less contracted(Nakai , 1915;Schneider , 1916).
Besides these two taxa , Nakai(1915)described the taxon with very short involucres and treat-
ed it as a new species , C.hallaisanensis based on Taquet s collections from Island Jeju_do of
South Korea , but Schneider(1916)designated it as C.sieboldiana var.brevirostris.The regional
flora (Lee , 1980;Lee , 1996)in Korea supported Nakai s circumscriptions and still maintained
this taxon at the specific level.On the other hand , C.sieboldiana var.mitis Nakai is character-
ized by its small fruits with a very narrow tube and a yellow_brownish pubescence mixed with a few
setose hairs (Schneider , 1916;Nakai , 1915), but has been currently recognized under C.
sieboldiana var.sieboldiana as one polymorphic species in Japan(Ohwi , 1984;Kitamura &Mura-
ta , 1984)due to the continuous variation of involucre width.
It is quite evident that a major difficulty in the taxonomy of the C.sieboldiana complex has
been the exclusive use of minor differences in f ruit morphology as well as leaf characteristics for
species definition and recognition.The confusion regarding the species delimitation was due to the
great variability of this complex from China to Japan throughKorea.There is a clear need for a study
of this complex that goes beyond the limited work that has been done previously as a small part of a
major floristic endeavor thus far.
The primary objective of this research was to redefine the phenetic relationships among the dif-
ferent morphological entities of the C.sieboldiana complex.Particular attention was focused on C.
mandshurica and C.sieboldiana in southern part of Korea which are quite different from C.
sieboldiana in Japan and C.mandshurica in northern part of Korea and northeastern China because
of the high degree of morphological intergradation among the entities and the inconsistent circum-
scription.It was also conducted to determine if the morphological differentiation among the C.
sieboldiana complex warrants specific taxonomic recognition with a large sample of field_collected
leaves as well as the pattern of geographical distribution.The effects of polymorphic level on leaf
shape and fruit size were investigated to determine the constancy of the characters.
No.3 CHANG Chin_Sung et al.:Multivariate morphometric study on Corylus sieboldiana complex 223　
1　Material and Methods
Specimens of Corylus sieboldiana var.sieboldiana and C.sieboldiana var.brevirostris were
obtained from University of Tokyo(TI), Tokyo Metropolitan University (MAK), Institute of Applied
Ecology , the Chinese Academy of Sciences , Shenyang(IFP), and Cheonnam University as loans as
well as the Arboretum of Seoul National University (SNUA).Mature leaves of C.mandshurica ,
which have been deposited at SNUA , were collected in many places in Korea and Japan from 1998
to 2002.Herbarium specimens of C.sieboldiana and C.mandshurica were selected to represent
the entire geographical range of Northeast China , Korea and Japan and to reflect the morphological
variability within each taxon.However , some specimens in Korea were almost indistinguishable mor-
phologically f rom both C.sieboldiana var.sieboldiana and C.mandshurica and these were re-
ferred to as the intermediate type for analysis.Therefore , two varieties of C.sieboldiana and C.
mandshurica were defined and analyzed here.Illustrations of the leaf and fruit morphologies(Fig.
1)were adapted from Nakai s drawing of C.sieboldiana (Nakai , 1915).
Fig.1.　The representative measured characters for the morphometric analysi s(from Nakai , 1915).
224　 Acta Phytotaxonomica Sinica Vol.42
Characters selected for analyses included those most frequently utilized in keys and diagnoses of
the various taxa.Characters considered useful by other authors(Schneider , 1916;Kitamura &Mu-
rata , 1984;Lee , 1980;Ohwi , 1984;Charkevicz , 1996)for reliable identification were also se-
lected.For leaf measurements a “ typical” and usually the largest measurable leaf was selected.The
initial data matrices were constructed from 18 characters(Table 1).Measurements were made with a
ruler and a binocular micrometer.One hundred and fifty two individuals of C.sieboldiana (74),
C.mandshurica (54), and the intermediate type (24)were measured for the various attributes.
All specimens used in the analyses were in a mature stage of development.
Criteria classically used in floras to distinguish between C.sieboldiana and C.mandshurica
concern differences in leaf shape and in involucre of the fruits(Kitamura &Murata , 1984;Ohwi ,
1984;Lee , 1980;Lee , 1996), but those traits do not clearly discriminate between the two species
because the variation of the traits is widely overlapped(see results).Therefore , based on the crite-
ria described by Schneider(1916), an individual of C.mandshurica observed in eastern Korea and
northeastern China represents the same characteristics as a plant of C.mandshurica in China and
Russia.Our approach consists of a study of individuals in northeastern China and eastern Korea and
individuals in Japan that we have classified , a priori , as C.mandshurica and C.sieboldiana re-
spectively .We then test a posteriori the capacity of measured characters to differentiate both groups
of individuals.Finally , we examine results from the only sites we found where the two species are
sympatric.Here we have found individuals(“ intermediate” plant)presenting some of the character-
istics of each species.Intermediate plants for the characters measured are of special importance in
testing the validity of our analysis.
Morphological variation within and among the taxa was assessed using univariate statistics
Table 1　Characters used in morphological analysis of C.sieboldiana complex＊
Leaf 1 Width
2 Length
3 Number of leaf veins
4 Number of serration in the apex
5 Length of apex
6 Angle of apex
7 Angle of base
8 Length between leaf veins
9 Length between leaf veins I(I)
10 Length between leaf veins II (J)
11 Length between leaf veins III(K)
17 Length of petiole
Fruit 12 Length of fruit
13 Number of serration of involucre
14 Length of serration of involucre
15 Width of lower part of involucre
16 Length of involucre(except nut)
18 Length of peduncle
＊ The numerals represent the same characters as shown in Fig.1.
No.3 CHANG Chin_Sung et al.:Multivariate morphometric study on Corylus sieboldiana complex 225　
(mean , maximum , minimum)and multivariate morphometric analyses(PCA)with SAS program (SAS
Institute Inc., 1988).A correlation matrix was generated using selected significant characters along with
analysis of variance (ANOVA)and univariate analyses.Also , bivariate analyses(bivariate scatter dia-
grams)were performed and each character associated with individuals of three OTUs(Operational Taxo-
nomic Units), C.mandshurica , C.sieboldiana and intermediate type , were plotted here.
2　Results
In principal components analysis , the first three factors accounted for 39.1%, 11.3%, and
9.4% of the total variance respectively , or 59.8%altogether.PC1 had the highest loadings for leaf
width(1)and length(2)and distance between lateral veins(9 to 11);PC2 had the highest load-
ings for fruit length(12), involucre length(16), serration depth of involucre(14), and width of
lower part of involucre(15), and PC3 had the highest loadings for number of lateral veins(3)and
peduncle length(18).PC1 provided more separation between C.sieboldiana and C.mandshurica
than PC2 and PC3 did.Leaf size and the distance between leaf veins were important characters that
permit the differentiation of these taxa.A plot on the second and third principal components , which
were related to characters of fruit , failed to establish discrete groups.Results of the PCA showed
that C.sieboldiana and C.mandshurica(Fig.2)were not morphologically distinct with morpho-
logical continuum.Indeed , the material of C.mandshurica in south Korea remains as a problematic
entity due to its morphological similarity to C.mandshurica and C.sieboldiana.
Univariate statistics , in addition to the minimum and maximum values for other characters
(Fig.3)showed that values overlapped extensively for these two taxa.The high correlation coeffi-
cients were found in several characters , such as , leaf length vs.width , and fruit length vs.involu-
cre length.
It is clear that C.sieboldiana and C.mandshurica are largely separated by the leaf length
and width (Schneider , 1916), i.e., the former elliptic , the latter oval except the intermediate
(Fig.4).If the intermediate individuals were included for the analysis , no strong discontinuities
exist between two taxa with respect to several diagnostic leaf characters.Much overlap between taxa
occurred in the central region of the scatter diagram as well as PCA(Figs.2 , 4).
Some individuals of C.mandshurica in south Korea are not greatly separated from the two
taxa , because they possess both the typical oval leaf of C.mandshurica and the elliptic leaf of C.
sieboldiana within the same twig for leaf morphology.
Besides the relationship between C.mandshurica and C.sieboldiana var.sieboldiana , C.
sieboldiana var.brevirostris and C.hallaisanensis , which have very short involucre , remained as
problematic taxa.The frequency distribution for involucre length(Fig.5)was examined in order to
test whether or not it was a diagnostic character for discrimination between taxa with short vs.long
involucre length.It revealed that distribution of the two species for this character overlapped consid-
erably more.Also , the separation of individuals with short length of involucre and those with inter-
mediate length of involucre on the distribution could not be considered entirely significant.There-
fore , there was virtually no separation of taxa with respect to the length of involucre tube.
3　Discussion
Corylus mandshurica in north and eastern Korea , northeastern China , and far eastern Russia
exhibited only the distinctive leaf shape and involucres(Ohwi , 1984;Schneider , 1916), compared
with C.sieboldiana.The extreme leaf form of C.mandshurica emerged as the most distinct mainly
due to the irregularly and coarsely serrate margin , and mucronate_acuminate or caudate apex , while
226　 Acta Phytotaxonomica Sinica Vol.42
Fig.2.　Scatter diagrams from principal components analysis of Corylus sieboldiana complex.　A , PC1 vs.PC2.B , PC1 vs.
PC3.◆, C.mandshurica;△, C.sieboldiana;＊, intermediate.
C.sieboldiana was characterized by its elliptic or oval leaves more or less gradually pointed at the
apex and regularly acute_serrate.In fact , Schneider (1916), who treated C.mandshurica as an
infraspecific taxon of C.sieboldiana , indicated that the extreme forms of both taxa looked very dis-
tinct.
Our initial examination of herbarium specimens suggested , however , the high degree of mor-
phological intergradation among its entities and its inconsistent circumscription within C.
sieboldiana complex.The morphometric analysis indicated that no strong discontinuities existed
among these taxa.Comparisons of the univariate , conceptually related means and range , pointed
up the similarity between C.sieboldiana and C.mandshurica.Therefore , the morphological
No.3 CHANG Chin_Sung et al.:Multivariate morphometric study on Corylus sieboldiana complex 227　
Fig.3.　Character variation of Corylus sieboldiana complex.　A , Number of leaf veins.B , Number of involucre serration.C ,
Depth of involucre serration.D , Width of tubular involucre.　I , intermediate(C.sieboldiana of southern Korea);M , C.mand-
shurica;S , C.sieboldiana.
Fig.4.　Scattered plot for leaf length and width of Corylus sieboldiana complex.　◆, C.mandshurica;□, C.sieboldiana;
＊, intermediate;C , China;K , Korea.
continuity among these taxa had prompted questions about their taxonomic status as species or vari-
eties.
Geographic distribution of the species should be taken into consideration along with the analysis
of similarities and differences of morphological characteristics as a basis for evaluating relationships
among taxa(Radford et al., 1974).The results obtained from this study do not support the separa-
tion of C.sieboldiana from C.mandshurica as an independent species.C.sieboldiana seemed to
be restricted to southern islands of Korea as well as Japan(Hokkaido to Kyushu)based on observa-
tion of herbarium specimens.C.mandshurica in southern parts of Korea was intermediate between
C.mandshurica from northern parts of Korea (Province Ganwg_won_do)and northeastern China
and C.sieboldiana from Japan and southern islands of Korea in leaf shape.It is shown from this
study that variation of the major key characters of C.sieboldiana in southern Korea was continuous
and overlapping with OTUs of C.mandshurica and C.sieboldiana from other areas.The
228　 Acta Phytotaxonomica Sinica Vol.42
Fig.5.　Frequency distribution of involucral bract length from Corylus sieboldiana var.sieboldiana (◆)and C.sieboldiana var.
mandshurica(□).
difficulties of quantitatively visualizing shape differences could be seen in the unclear distinction
among OTUs in southern Korea.Therefore , it exhibited recombination of some traits , resembling
both C.sieboldiana and C.mandshurica in leaf serration , leaf apex , and short involucre tube.
Corylus sieboldiana var.brevirostris , which has been recognized as morphologically distinct
from C.sieboldiana var.sieboldiana in Japan , remains as a problematic taxon (Schneider ,
1916).A detailed investigation of many individuals of C.sieboldiana in Japan revealed a compara-
ble degree of continuous variation of bract length and width.Many specimens of short involucre type
(=C.sieboldiana var.brevirostris)exhibited widely overlapping ranges in the length of involucre
with C.sieboldiana var.sieboldiana (Fig.2).The frequencies for involucre length (Fig.3)
showed the normal distribution , suggesting this characterwas related to the polygenic trait.Both ob-
served intrapopulation variation and the presumed simple genetic control of the character would sug-
gest that it is insufficient for recognition of species or infraspecific taxon.Although extreme entity
was specific to some individuals , none of these individuals could be reliably , consistently , and geo-
graphically identified with the use of this character alone.Therefore , the previously recognized
taxon , C.sieboldiana var.brevirostris , should be united with C.sieboldiana along with forms in-
termediate in fruit and involucre morphology between these two taxa.C.hallaisanensis , which was
described by Nakai(1915)in southern Korea based on short involucre , has never been compared
with C.sieboldiana var.brevirostris in Japan thus far.When Schneider(1916)described the short
involucre type of C.sieboldiana as an infraspecific species based on the Japanese individuals , he
was not able to examine Nakai s type specimen (collected from Island Jeju_do of Korea , Nakai ,
1915).Therefore , Schneider (1916)never compared the major difference between C.sieboldiana
var.brevistoris and C.hallaisanensis.Unfortunately , we were not able to confirm Nakai s type
specimen on C.hallaisanensis at TI either and only usedNakai s drawing (Fig.2 of Flora Sylvati-
ca Koreana , Nakai , 1915).This study demonstrated that taxa of Corylus sieboldiana complex can-
not be delimited with the use of leaf morphological characters alone.Therefore , a plausible sugges-
tion is that the entities of this group (=C.hallaisanensis and C.sieboldiana var.brevirostris)
should be united and recognized under one taxon , C.sieboldiana var.sieboldiana.
No.3 CHANG Chin_Sung et al.:Multivariate morphometric study on Corylus sieboldiana complex 229　
Fig.6.　A map of Corylus sieboldiana var.sieboldiana(◆)
and C.sieboldiana var.mandshurica (★, Chung , 1943;
○, based on specimens of SNUA)in Korea.
　　According to Li and Cheng (1979), C.
sieboldiana complex as well as C.chinensis
Franch.and C.fargesii Schneid.were all mem-
bers of sect.Corylus , subsect.Siphonochlamys.
However , Whitcher and Wen(2001), Forest and
Bruneau(2000), and Erdogan and Mehlenbacher
(2000) show that C.chinensis , which has
deeply dissected involucral margins and a tree
habit , is phylogenetically different from C.
sieboldiana complex with respect of ITS phyloge-
ny .On the other hand , the monophyly of C.
mandshurica and C.sieboldiana is strongly sup-
ported by the ITS phylogeny (Forest &Bruneau ,
2000).Therefore , C.mandshurica and C.
sieboldiana may be of common origin.
The entities that comprise C.sieboldiana
complex exhibit widely overlapping ranges in all
morphological attributes.Consequently , each en-
tity cannot be considered to be consistently dis-
tinct.The most cohesive vegetative morphology is
largely due to the high level of variability ex-
pressed in the leaves.The leaf and fruit data may
reflect a lack of divergence between them.We do
not believe these to be sufficient for taxonomic
splitting of C.mandshurica from C.sieboldiana
to warrant the designation of the rank of a species
based on morphology and their origin.The two
taxawere not morphologically well differentiated where their ranges of distribution come together
(Fig.6).
In summary , based on our results from the morphometric analyses , C.sieboldiana complex is
interpreted as one species , C.sieboldiana , with one infraspecific taxon , C.sieboldiana var.
mandshurica .Therefore , our systematic interpretation on C.sieboldiana complex was in agreement
with the taxonomic treatment of Schneider (1916)except C.sieboldiana var.brevistoris.This
study confirmed that C.sieboldiana var.sieboldiana is distributed in southern Korea as well as in
Japan , while C.sieboldiana var.mandshurica is found in Korea , northeastern China , and far
eastern Russia.Although C.sieboldiana var.mandshurica in Japan has been recorded to occur
from Hokkaido (Ohwi , 1984;Kitamura &Murata , 1984 , Figs.6 , 7), no material was seen from
MAK and TI this time.This locality was not included on the map of distribution of C.sieboldiana
var.mandshurica.Based on our analyses , the following key and taxonomic treatments are provided
as follows.
Key to the taxa of Corylus sieboldiana complex
1.Leaves elliptic or oval , margin regularly acute_serrate , apex more or less gradually pointed;lateral veins(7)8-
9(12)on each side of midvein;tubular involucre shallowly divided into lobes , lobes(0)2-6(8)mm long;nar-
rower tube of involucre 2-8 mm wide C.sieboldiana var.sieboldiana………………………………………
1.Leaves orbicular_ovate , margin irregularly and coarsely serrate , apex mucronate_acuminate or caudate;lateral
230　 Acta Phytotaxonomica Sinica Vol.42
veins(6)7-8(10)on each side of midvein;tubular involucre more deeply divided into linear lobes , lobes(3)5
-8(14)mm long;narrower tube of involucre 3-12 mm wide C.sieboldiana var.mandshurica…………
Fig.7.　Distributions of Corylus sieboldiana var.mandshurica and C.sieboldiana var.sieboldiana of eastern Asia.　Adapted
f rom Charkevicz , 1996;Fu et al., 2000;Ohwi , 1984.
4　Taxonomic treatments of Corylus sieboldiana complex
Corylus sieboldiana Blume in Ann.Mus.Bot.Ludg._Bat.1:310.1850.——— Corylus
heterophylla Fisch.ex Trautv.var.sieboldiana (Blume)A.DC., Prodr.Systematis Naturalis
Regni Vegetabilis 16 , 2:113.1864.——— Corylus rostrata Aiton var.sieboldiana (Blume)Max-
im.in Bull.Acad.Imp.Sci.Saint_Pétersbourg 27:538.1882.Type:unknown.
Corylus rostrata Aiton var.mitis Maxim.in Bull.Acad.Imp.Sci.Saint_Pétersbourg 27:
538.1882.——— Corylus sieboldiana Blume var.mitis (Maxim.)Nakai in Bot.Mag.(Tokyo)
29:37.1915.Type:Japan.Suruga , Mt.Fuji san , 1864.S.Tschonoski s.n.(isotype , A !,
seen as a photo).
Corylus sieboldiana var.brevirostris C.K.Schneid.in Sargent , Plantae Wilsonianae 2:453.
1916.——— Corylus brevirostris(C.K.Schneid.)Miyabe , J.Fac.Agric.Hokkaido Univ.26:
458.1934.Type:Japan.Hokkaido , O_shima , Shiribeshi San , 1914_08_27.E.H.Wilson 7268
(holotype , A !, seen as a photo).
Corylus hallaisanensis Nakai , Feddes Repert.13:350.1914.
Type:Korea.Querlpaert(=Island Jeju_do), Hallai_san(=Mt.Halla_san), forest , 1903_
09 , Taquet 333(isotype , TI!).
var.sieboldiana
Distribution:Japan(Hokkaido to Kyushu), Korea (southern islands)
Representative specimens:Japan.Aichi:Kitasitara_gun , Toyone_mura , Chausu_yama , H.
T.Im 3142_1(Cheonnam National University).Aomori:Mt.Hakkoda , between Shimokenashitai
No.3 CHANG Chin_Sung et al.:Multivariate morphometric study on Corylus sieboldiana complex 231　
Moor and Sukayu Spa , Koji Yonekura 4265 (MAK);Nakatsugaru_gun , Iwaki_cho , H.T.Im
8243_1 (Cheonnam National University);Aomori_shi , Mt.Hakida_san , C.S.Chang 885
(SNUA).Gifu:Gujyo_gun , Takemura , Hirugano , G.Murata 14432 (MAK).Gunma:Agat-
suma_gun , Takayaka Valley , T.Kawahara &M.Maki 5768 (TI).Hokkaido:Soraichi_sicho ,
Asibechu_si , Kirigisu_yama , H.T.Im &J.Murata 23125_1 (Cheonnam National University).
Kyoto:Kitakuwada_gun , Chii_mura , Ashiu , Nakayama , S.Okamoto s.n.(MAK).Miyazaki:
Higashiusuki_gun , Shiiba_mura , M.Hotta 6574 (MAK);Nishiusuki_gun , Gogasa_cho , Shirai-
watoge , Mt.Shiraiwa , H.T.Im 2865_1(Cheonnam National University).Nagano:Mt.Hachin-
buse , S.Momose s.n.(TI);Shimo_takai_gun , Tamanouchi_machi , Shiga Heights , Near Biwa
Pond , M.Mizushima s.n.(MAK);Yamanakaku_mura , T.Togashi s.n.(MAK).Okayama:
from Mt.Maruyama , 13 km to Mt.Daisen , Kawakami_son , Hidenobu Funakoshi 1348_1(Cheon-
nam National University).Shiga:Ika_gun , Yogo_cho , H.T.Im 10510 (TI).Tochigi:Nikko ,
near Jakko_no_taki , J.Murata &H.Ohashi s.n.(TI).Tokyo:Nishi_tama_gun , Okutama_
machi , Nippara , Mt.Tenso_zan , M.Mizushima s.n.(MAK);Hachioji_shi , from Katakura_cho
and Kobiki_cho , H.T.Im &T.Kawahara 10462(TI).Yamanashi:Mt.Fuji , Shuchi Noshiro
2759(TI);Minamikoma_gun , Minobu_cho , Mt.Tenshigatake , Y.Kadota & J.Murata 3150
(TI);Minamisturu_gun , Mt.Mistutouge_yama , C.S.Chang 1398 (SNUA);Minamitsuru_gun ,
Kawakuchiko_cho , H.T.Im &C.S.Park 33939(Cheonnam National University);Enzan_shi ,
H.T.Im 4750(Cheonnam National University).
Korea.Jeollabuk_do:Mt.Baek_yang_san , T.Lee et al.6290 (SNUA);Mt.Nae_jang_
san , T.Lee et al.6053(SNUA), C.S.Chang 3657(SNUA).Jeollanam_do:Gurye_gun , Mt.
Ji_ri_san , Nogo_dan , T.Lee &C.S.Chang s.n.(SNUA);Mt.Jo_gae_san , T.Lee s.n.
(SNUA);Island Hong_do , T.Lee s.n.(SNUA);Wando_gun , Island Wan_do , B.Yinger et al.
3449(SNUA);Jaseong_gun , C.S.Chang et al.3657(SNUA);Gwang_yang_si , T.Lee s.n.
(SNUA);Hwasun_gun , H.T.Im &C.G.Yang 21726(Cheonnam National University);Sinan_
gun , Zi_do , H.T.Im et al.38363(Cheonnam National University);Jangheung_gun , H.T .Im
34715(Cheonnam National University);Gwangju , Dong_gu , Chiwon_dong , H.T.Im &C.S.
Park 33881(Cheonnam National University).Gyonggi_do:Yangpyeong_gun , Yong_mun_san , T.
Lee s.n.(SNUA);Ong_jin_gun , Island Paekyong_do , B.Yinger et al.2248(SNUA);Uiwang_
si , H.S.Oh &B.S.Mun s.n.(SNUA).Cheju_do:Cheju_shi , Odung_dong , J.Murata et al.
21054(Cheonnam National University).
var.mandshurica(Maxim.&Rupr.)Schneid.in Sargent , Plantae Wilsonianae 2:454.
1916.——— Corylus mandshurica Maxim.&Rupr., Bull.Acad.Imp.Sci.Saint_Pétersbourg 15:
137.1856.——— Corylus rostrata var.mandshurica(Maxim.&Rupr.)Regel , Bull.Acad.Imp.
Sci.Saint_Pétersbourg 15:221.1857.Type:Russia.Amur river , no date , C.J.Maximowicz
s.n.(A !, seen as a photo).
Distribution:China , Russia , Korea , and Japan(Hokkaido).
Representative specimens:Korea.Chungcheongbuk_do: Jecheon_si , T.Lee s.n.
(SNUA).Chungcheongnam_do:Mt.Goe_rong_san , T.Lee et al.s.n.(SNUA);Yesan_gun ,
Mt.Ga_ya_san , C.S.Chang s.n.(SNUA).Gangwan_do:Myongju_gun , H.T.Im s.n.
(Cheonnam National University);Yangyang_gun.Osek_oncheon , Mt.Sul_ak_san , J.Murata &S.
C.Ko 21072(Cheonnam National University);Jeongsun_gun , H.T.Im 34919_1(Cheonnam Na-
tional University);Wonju_si , Socho_myeon , Mt.Chi_ak_san , C.S.Chang &J.I.Jeon 1884
(SNUA);Inje_gun , Mt.Jumbong_san , C.S.Chang et al.3629(SNUA);Yanggu_gun and Inje_
gun , T.Lee et al.s.n.(SNUA);Yanggu_gun , C.S.Chang et al.BS052(SNUA);Yeongwol_
232　 Acta Phytotaxonomica Sinica Vol.42
gun , Jeon &H.S.Lee 11000(SNUA);Goseong_gun , C.S.Chang et al.HR255 (SNUA);
Hwacheon_gun , T.Lee s.n.(SNUA);Pyeongchang_gun , Dae_Gwan_ryeong , T .Lee &M.Y.
Cho s.n.(SNUA);Pyeongchang_gun , C.S.Chang &J.I.Jeon 1984(SNUA);Pyeongchang_
gun , C.S.Chang &Choi 1608 (SNUA);Taebak_si , C.S.Chang &Kim 1904 (SNUA).
Gyeongsangbuk_do:Mungyeong_si , Mungyeong_eup , C.S.Chang et al.UD005 (SNUA);
Mungyeong_si , Mungyeong_eup , Jo_Ryeong , C.S.Chang & J.I.Jeon 1958 (SNUA);
Mungyeong_si , Mungyeong_eup , T.Lee s.n.(SNUA);Mungyeong_si , C.S.Chang &Kim
OJ019(SNUA);Uljin_gun , D.R.Choi &S.M.Jang BA095 (SNUA);Yeongyang_gun , Ju_
san , D.R.Choi &S.G.Kwon JOO081(SNUA).Gyeonggi_do:Pocheon_gun , Gwang_neung ,
T.Lee s.n.(SNUA);Anyang_si , T.Lee s.n.(SNUA);Inchon , Ganghwa_gun , T.Lee s.n.
(SNUA);Yangpyeong_gun , C.S.Chang &J.I.Jeon 1831(SNUA);Suwon_si , T.Lee s.n.
(SNUA);Gapyeong_gun , C.S.Chang &J.I.Jeon 1904 (SNUA).Jeollanam_do:Jangsung_
gun , H.T.Im 22218(Cheonnam National University).
China.Heilongjiang:Yichun , T.Y.Ding et al.2433(IFP);Acheng , G.Z.Wang et al.
444 , W.Wang et al.785(IFP).Jilin:Antu , T.N.Liou 3862(IFP);Changbai , S.D.Zhao
&S.Q.Zhong 2472(IFP), S.X.Li et al.733 , 1005(IFP);Fusong , T.N.Liou et al.1271
(IFP);Hunjiang (Linjiang), S.X.Li et al.1155 (IFP);Wangqin , P.Y.Fu et al.862
(IFP).Liaoning:Benxi , W.Wang et al.409 (IFP);Huanren , C.S.Wang 4126 (IFP), J.
Y.Li et al.1907(IFP), S.C.Chang &Y.C.Zhu 214(IFP), Y.C.Deng &K.Z .Wang
671(IFP);Lingyuan , Exped.Team 178(IFP);Qingyuan , C.S.Wang et al.1210(IFP).
Acknowledgements　We thank Ms.Woo_Kyong Ahn for her assistance in the measurement of
specimens.We would like to thank Drs.Akiko Shimizu at TI , Machio Wakabayashi at MAK ,
Hyung_Tak Im at Cheonnam National University , and Cao Wei at Institute of Applied Ecology , the
Chinese Academy of Sciences , Shenyang (IFP)for loaning their collections of Corylus.We also
thank the anonymous reviewer for providing helpful comments.
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Appendix.Origin and accession number for specimens utilized for morphological analysis.
C.sieboldiana var.sieboldiana
Japan.Aomori:C.S.Chang et al.885 (SNUA);H.T.Im 8243_1 , 8243_2 , 10574 , 23211(Cheonnam
National University);Mt.Hakkoda , between Shimokenashitai Moor and Sukayu Spa , K.Yonekura s.n.(MAK).
Aichi:H.T.Im &J.Murata 3142 , 3142_1 , 3142_3 , 3142_5 (Cheonnam National University).Akita:Tsubaki ,
Oga Peninsula , N.Satomi s.n.(MAK).Fukushima:Kashi , K.Sugawara s.n.(MAK).Gifu:Hirugano , Taka-
su_mura , Gujyo_gun , Mino , Hondo , G.Murata s.n.(MAK).Gunma:Kawahara&Maki 5768(TI);J.Murata &
H.Ohashi s.n.(TI);Kyu_shikazawa , T.Yano s.n.(MAK).Ishikawa:Sue , Mii , Wajima City , N.Satomi s.n.
(MAK).Kyoto:Nakayama in Ashiu , Chii_mura , Kitakuwada_gun , S.Okamoto s.n.(MAK).Miyagi:Mt.
Kurikoma(Kurikoma Town), H.Ishikawa s.n.(MAK).Miyazaki:H.T.Im 2865_1 , 2865_2(Cheonnam Nation-
al University);en route from Kubikukuri to Goyozan , Shiiba_mura, Higashiusuki_gun , M.Hotta s.n.(MAK).
Nagano:Gofukuji , Matsumoto City , H.Okuhara s.n.(MAK);Tanaba , Omachi City , N.Kato s.n.(MAK);
Lake Nojiri(Shinano Town), T.Yano s.n.(MAK);Near Biwa Pond , Shiga Heights , Yamanouchi_machi , Shimo_
takai_gun , M.Mizushima s.n.(MAK);Yanaba , N.Kato s.n.(MAK).Okayama:Funakoshi 1348_1 , 1348_2
(Cheonnam National University).Shiga:H.T.Im 10510 , 10510_1 , 10510_2(Cheonnam National University);H.
T.Im & J.Murata 4661 (Cheonnam National University).Tochigi:Nikko , Jakko , Takimichi , M.Ono &S.Ono
s.n.(MAK).Shimotsuke , Nikko(Nikko City), T.Makino s.n.(MAK).Tokyo:H.T.Im &Kawahara 10462
(TI);Nippara_Mt.Tenso_zan , Okutama_machi , Nishitama_gun , M.Mizushima s.n.(MAK).Tottori:Mt.
Daisen , N.Satomi s.n.(MAK);Mt.Kawanori , Okutama_machi , K.Suzuki s.n.(MAK).Yamanashi:C.S.
Chang &H.T.Im.1398 (SNUA);N.Shuichi 2759(TI);Tateishi&J.Murata 3150(TI);Yamanakako_mura ,
M.Togashi s.n.(MAK);Okura_takamaru , Yamato Village , Higashi_yamanashi Co., M.Tsuchiy a s.n.(MAK);
Mitsutoge Pass , Kai , K.Hiyama s.n.(MAK);H.T.Im &Park 33939(Cheonnam National University).Hokkai-
do:H.T.Im & J.Murata 4581 , 4581_1 , 4581_2 , 23125_1 , 23125_2 , 23125_3(Cheonnam National University).
Korea.Gyeonggi_do:B.R.Yinger et al.2248(SNUA).Jeollanam_do:T.Lee s.n.(SNUA);R.Dudley
&B.R.Yinger 3449 (SNUA);H.T.Im 7335 , 22218 , 34715 , 38363(Cheonnam National University).
C.sieboldiana var.mandshurica (=C.mandshurica)
Korea.Gyeonggi_do:C.S.Chang &J.I.Jeon 1904(SNUA);T.Lee s.n.(SNUA).Gangwon_do:C.
S.Chang &S.Y.Choi 1608(SNUA);J.I.Jeon &H.S.Lee 10973(SNUA);C.S.Chang 3629(SNUA);C.
S.Chang &J.I.Jeon 1884(SNUA);C.S.Chang &H.Kim 3016 , 3021 (SNUA);C.S.Chang et al.3644 ,
HR 255 , PA026 , SM 051 , SM 052(SNUA);T.Lee s.n.(SNUA 7341), T.Lee s.n.(SNUA 7318), T.Lee s.
n.(SNUA 7340), T.Lee s.n.(SNUA 7368), T.Lee s.n.(SNUA 7276), T.Lee&M.Y.Cho s.n.(SNUA),
J.Murata &S.C.Ko 21072 (Cheonnam National University), H.T.Im 34919_1 , 35008(Cheonnam National
234　 Acta Phytotaxonomica Sinica Vol.42
University).
China.Heilongjiang:T.Y.Ding et al.2433 (IFP), G.Z.Wang et al.444(IFP), W.Wang et al.785
(IFP).Liaoning:J.Y.Li et al.1907 (IFP), S.C.Cui &Y.C.Zhu 214 (IFP), C.S.Wang et al.1210
(IFP), C.S.Wang 4126(IFP), W.Wang et al.409(IFP), Heilongjiang Exped.Team 178(IFP), Y.C.Deng
&K.Z.Wang 671(IFP).Jilin:S.X.Li et al.1155(IFP), T.N.Liou et al.1271(IFP), S.X.Li et al.733
(IFP), S.D.Zhao &Z.S.Qin 2472(IFP), P.Y.Fu et al.862(IFP), T.N.Liou 3862(IFP), S.X.Li et
al.1005(IFP).
Intermediate form(=C.sieboldiana var.sieboldiana)
Korea.Gyeonggi_do:C.S.Chang&J.I.Jeon 1831(SNUA), T.Lee&C.S.Chang s.n.(SNUA), T.
Lee s.n.(SNUA), T.Lee , s.n.(SNUA), T.Lee et al.s.n.(SNUA), T.Lee s.n.(SNUA).Gyeongsang-
buk_do:C.S.Chang &J.I.Jeon 1958 (SNUA), C.S.Chang s.n.(SNUA).Jeollabuk_do:T.Lee s.n.
(SNUA), T.Lee s.n.(SNUA), T.Lee &C.S.Chang s.n.(SNUA).Jeollanam_do:T.Lee s.n.(SNUA),
C.S.Chang 2982(SNUA), T.Lee s.n.(SNUA), T.Lee &C.S.Chang s.n.(SNUA).Chungcheongnam_
do:T.Lee &M.Y.Cho s.n.(SNUA).
毛榛复合体(桦木科)多变量形态学研究及分类学处理
1
CHANG Chin_Sung 　1CHANG Gae_Sun 　2覃 海 宁
1(汉城国立大学森林资源系暨树木园　水原 441_744　韩国)
2(中国科学院植物研究所　北京 100093　中国)
摘要　Corylus mandshurica Maxim.&Rupr.和 C.sieboldiana Blume在中国和俄罗斯一直被处理为独立的
种。本文对来自不同标本室的这两个种的腊叶标本及野外采集的大量叶片进行了形态学分析 , 以确定
形态分化是否足够成为建立种的依据。对采自中国 、韩国和日本覆盖两个种分布区的 153 份标本的 18
个形态学性状进行了统计 , 以构建数据矩阵用于主成分分析。结果表明 ,包含 C.sieboldiana 复合体的
数据单位(entities)在所有的形态学性状上均呈现出广泛的重叠区域。叶片和果实性状分析结果表明类
群间缺乏分异性。基于形态学性状不足以把 C.mandshurica 从C.sieboldiana 中分离出去 , 并给予种的
等级 , 也就是说这两个类群在形态上没有完全分化 ,而且它们的分布区也是相连的。因此 , 应该把 C.
mandshurica 作为C.sieboldiana 的种下分类群处理。 C.sieboldiana 在韩国南部和日本的具短总苞的植物
常常被处理为独立的种或变种 ,但实际上应该作为 C.sieboldiana 内的变型处理 , 因为总苞性状高度变
异 ,即使在同一个体上也是如此。
关键词　毛榛复合体;形态度量主成分分析;分类学处理
No.3 CHANG Chin_Sung et al.:Multivariate morphometric study on Corylus sieboldiana complex 235