Taxonomy and phylogeography of Cardamine impatiens …ibot.sav.sk/usr/Karol/docs/PDF_files/Kucera et al. BJLS_2006.pdf · Taxonomy and phylogeography of Cardamine impatiens ... did

Botanical Journal of the Linnean Society

2006

152

169ndash195 With 7 figures

copy 2006 The Linnean Society of London


2006

152

169ndash195

169

Blackwell Publishing LtdOxford UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London 2006 20061522169195Original Article

TAXONOMY OF C IMPATIENS AND C PECTINATAJ KU

B

ERA ET AL

Corresponding author E-mail karolmarholdsavbask

Taxonomy and phylogeography of

Cardamine impatiens

and

C pectinata

(Brassicaceae)

JAROMIacuteR KU

B

ERA

1

JUDITA LIHOVAacute

1

and KAROL MARHOLD

FLS12

1

Institute of Botany Slovak Academy of Sciences Duacutebravskaacute cesta 14 SK-845 23 Bratislava Slovak Republic

2

Department of Botany Charles University Benaacutetskaacute 2 CZ-128 01 Praha 2 Czech Republic

Received November 2005 accepted for publication March 2006

The taxonomic position of

Cardamine pectinata

a controversial taxon treated either as infraspecific to

C impatiens

or as a separate species was studied Forty-nine populations were sampled and used in multivariate morphometricand molecular (amplified fragment length polymorphism) analyses Our results showed that

C impatiens

and

C pectinata

represent two well-differentiated taxa in terms of both molecular and morphological data and should betreated as two separate species We present the taxonomic conspectus for both species including the list of synonymsinformation on type specimens morphological descriptions ecological characteristics and distribution area Typespecimens for several names are designated here The detailed geographical distribution of

C pectinata

is presentedbased on the survey and revision of herbarium material It covers the area from the Balkans through Turkey and theCaucasus to Iran As an introduction to the phylogeography of both species the distribution of genetic diversitywithin and between the studied populations and geographical regions was assessed In

C pectinata

the highestgenetic diversity was observed in northern Turkey suggesting the location of glacial refugia along the Black Seacoast Populations in Bulgaria were less diverse and this area has most probably been colonized postglacially

C impatiens

on the other hand did not display clear phylogeographical structure most probably as a result of theefficient spread and mixing between different colonization routes copy 2006 The Linnean Society of London

BotanicalJournal of the Linnean Society

2006

152

169ndash195

ADDITIONAL KEYWORDS

amplified fragment length polymorphism ndash Balkan ndash Caucasus ndash chromosomenumbers ndash Cruciferae ndash genetic diversity ndash glacial refugia ndash multivariate morphometrics ndash nomenclature ndash

Turkey

INTRODUCTION

The genus

Cardamine

L (Brassicaceae) comprisesabout 200 species occurring indigenously on all conti-nents except mainland Antarctica (Al-Shehbaz 1988)Diploid as well as polyploid taxa up to high ploidy levelsare represented (reviewed by Lihovaacute amp Marhold 2006)Although several detailed studies have been publishedon polyploid complexes of

Cardamine

in Europe(

C pratensis

group Loumlvkvist 1956 Marhold 1994aUrbanska

et al

1997 Franzke amp Hurka 2000 LihovaacuteMarhold amp Tribsch 2003 Lihovaacute Tribsch amp Stuessy2004b

C amara

group Loumlvkvist 1957 Marhold 19921995 Lihovaacute Marhold amp Neuffer 2000 Lihovaacute

et al

2004a

C raphanifolia

group Pern

y

et al

2005a) less

attention has been paid so far to differentiation pat-terns in diploids Morphometric and molecular analy-ses of the Balkan diploid

C acris

have recently led tothe recognition of three subspecies within this area(Pern

y

Tribsch amp Anchev 2005b) Variation patterns inthe diploid populations of the widespread

C amara

reflect a rather recent isolation and divergence of pop-ulations restricted to different European mountainranges which are now classified as subspecies (Lihovaacute

et al

2000 2004a Marhold Huthmann amp Hurka2002) On the other hand ongoing studies on othergroups of diploid taxa the ItalianndashBalkan

C maritima

(J Ku

d

era J Lihovaacute amp K Marhold unpubl data) and

C glauca

(Laku

t

i

c

et al

2006) indicate much morepronounced diversification in terms of both morphologyand molecular markers than revealed in the above-mentioned taxa and species groups

170

J KU

B

ERA

ET AL



2006

152

169ndash195

Cardamine impatiens

L is a diploid species with awide distribution in Eurasia (Jalas amp Suominen1994) reported as introduced also from North America(Seymour 1969 Rollins 1993) and southern Africa(Dreyer amp Jordaan 2000) Although several infraspe-cific taxa have been described in the past within thisspecies most of them appear currently only in synon-ymy The sole exception is the taxon originallydescribed as

C pectinata

Pall ex DC by Candolle(1821 264) based on the material collected by Pallasin Persia (today Iran) Apart from Iran (Hedge 1968)it is also reported from the Caucasus Turkey and theBalkan Peninsula (Stoyanov amp Stefanov 1948 Gross-geim 1950 Cullen 1965 Assenov 1970 Ketskhoveli1979 Kit Tan 2002) The taxonomic status of thistaxon however has remained controversial It hasbeen treated either as a separate species (by Gross-geim 1950 Assenov 1970 Ketskhoveli 1979) or as asubspecies (by Stoyanov amp Stefanov 1948 Jones ampAkeroyd 1993) or variety (by Trautvetter 1876Zohary Heyn amp Heller 1980) of

C impatiens

Cullen(1965 443) treating this taxon at the varietal rankargued that lsquointermediates between the two varieties[var

impatiens

and var

pectinata

] are of frequentoccurrencersquo This opinion has been shared also by KitTan (2002 184) and Khatri (1988) although the latterauthor treated

C pectinata

at the species level Otherauthors (eg Assenov 1970) have no doubt that thisshould be a clearly separated species

Although

C impatiens

exhibits a wide distributionacross Eurasia the area of the present occurrence of

C pectinata

is regarded to be restricted to the BalkanPeninsula Turkey Iran and lower altitudes of theCaucasus (see references above) These areas areknown to harbour considerable taxonomic and geneticdiversity that can be attributed to processes associ-ated with Pleistocene glaciations Palaeobotanical andphylogeographical studies have shown that the south-ern peninsulas of Iberia Apennines Balkans and theCaspianCaucasus region served as refugia in whichmany European temperate species survived unfavour-able climatic conditions during the glacial periods(Hewitt 2004) Repeated cycles of area contractionand expansion as well as altitudinal shifts in south-ern mountain ranges allowed considerable geneticdiversity to be retained in these regions in contrastwith the genetic impoverishment often observed inmore northern areas (Hewitt 2002 2004) Severalcontinental-scale phylogeographies documentingthese processes have been published in recent yearsaddressing mainly tree species (eg King amp Ferris1998 Fineschi

et al

2002 Hampe

et al

2003Heuertz

et al

2004) A few phylogeographical studiesalso covered or at least marginally included theeasternmost putative refugial areas in Turkey andthe CaspianCaucasus region (King amp Ferris 1998

Hampe

et al

2003 Marcussen 2003 Albach Mar-tiacutenez-Ortega amp Chase 2004) Reconstructions of thevegetation in the last glacial maximum have docu-mented the occurrence of broad-leaved deciduous andmixed forests in the area along the Black Sea coast ofTurkey and Georgia (Adams 1997) This region char-acterized by many Tertiary and Quaternary relic spe-cies (Denk Frotzler amp Davitashvilli 2001) may haveprovided favourable habitats for the survival of sev-eral temperate herbaceous species including the

Car-damine

taxa studied hereTwo main questions are addressed in this study

First we aimed to resolve the taxonomic position ofpopulations assigned to

C pectinata

in respect of thewidespread

C impatiens

Detailed morphometric andmolecular (amplified fragment length polymorphismAFLP) analyses have been carried out together withthe survey of herbarium records from the sympatricregions Second as an introduction to the phylogeog-raphy of both species the distribution of genetic diver-sity in the studied populations and geographicalregions was assessed using AFLP data

MATERIAL AND METHODS

P

LANT

MATERIAL

Population sampling was focused on the area with thereported occurrence of both taxa

C impatiens

and

C pectinata

ie the Balkan Peninsula northern Tur-key and the Caucasus region

C impatiens

was alsosampled from numerous populations in more distantparts of its distribution range (central and southernEurope Japan) to cover the area without the occur-rence of

C pectinata

(Table 1 Fig 1) For morphomet-ric analyses herbarium specimens of a total of 565individuals originating from 26 populations were used(6ndash34 plants per population) These were collected inthe sympatric area of the two taxa Material for AFLPanalyses (silica gel-dried leaves) was represented by82 samples from 24 localities also sampled for mor-phometric studies and 65 samples from an additional23 populations collected mainly in more remote geo-graphical areas (Table 1) For chromosome countsseeds originating from 15 localities were used Thesewere either collected directly in the field or fromplants collected at the localities and transplanted inthe experimental field in Bratislava Slovak Republic

Details on the origins of the plant material are givenin Table 1 Voucher specimens are deposited in theherbarium SAV

To present a detailed view on the distribution ofplants corresponding to

C impatiens

and

C pectinata

especially in the area of their sympatric occurrenceherbarium material from the following herbaria wasrevised B BM G GH K KW LE LEU P PR PRC

TAXONOMY OF

C IMPATIENS

AND

C PECTINATA

171



2006

152

169ndash195

Table 1

List of populations of

Cardamine impatiens

and

C pectinata

used in karyological (2

n

) morphometric (Morph)and amplified fragment length polymorphism (AFLP) analyses (ndash not used in the particular analysis) Data on chromosomenumbers represent new reports Note that chromosome numbers were counted also from two Korean populations of

C impatiens

not used in morphological or AFLP analyses Voucher specimens are deposited in SAV Collectors

HK

HKato

HKu

H Kudoh

JK

J Ku

d

era

JL

J Lihovaacute

KM

K Marhold

MK

M Kolniacutek

MO

M Onette

MS

M Slovaacutek

SK

S Kato

SS

S Stoyanov

VGO

V G Onipchenko

Population code origin and collection data 2

n

Morphno ind

AFLP no ind

Cardamine impatiens

LI-Bu-1 ndash Bulgaria Plana Mts near village Dolny Pasarel valley of Iskar river

42

deg

32

prime

35

Prime

N 23

deg

29

prime

02

Prime

E 660 m 1vi2003

JK amp SS

16 30 1

I-Bu-2 ndash Bulgaria Plana Mts near lake Pasarel under Mt Asanitsa 43

deg32prime59PrimeN 23deg28prime17PrimeE 710 m 1vi2003 JK amp SS

ndash 30 3

I-Bu-3 ndash Bulgaria Stara planina Mts near village Kalofer along the river Tundya 42deg37prime31PrimeN 24deg58prime00PrimeE 680 m 2vi2003 JK amp SS

ndash 7 3

I-Bu-4 ndash Bulgaria Stara planina Mts Kalofer Panitsite 42deg39prime25PrimeN 24deg58prime51PrimeE 740 m 2vi2003 JK amp SS

ndash 24 3

I-Bu-5 ndash Bulgaria Vasiliovska planina Mts c 3 km W of Bogoya (saddle) stream Kozidol 42deg51prime39PrimeN 24deg28prime43PrimeE 910 m 4vi2003 JK amp SS

ndash 30 1

I-Bu-6 ndash Bulgaria between Vasiliovska planina Mts and Tetevensky Balkan Mts c 5 km S of village Ribaritsa near spring Borisova cheshma valley of Debelovitsa river 42deg50prime54PrimeN 24deg26prime45PrimeE 780 m 4vi2003 JK amp SS

ndash 8 2

I-Bu-7 ndash Bulgaria Tetevenska planina Mts valley of Vidima river near village Ribaritsa 42deg45prime05PrimeN 24deg54prime20PrimeE 1010 m 5vi2003 JK amp SS

16 29 3

I-Bu-8 ndash Bulgaria Tetevenska planina Mts valley of Zavodna river along the road from Ribaritsa to cottage Vezhen c 2 km S of Ribaritsa village 42deg48prime48PrimeN 24deg22prime17PrimeE 760 m 5vi2003 JK amp SS

16 25 3

I-Bu-9 ndash Bulgaria Tetevenska planina Mts under the cottage Vezhen near the tourist path from valley of Zavodna river 42deg45prime35PrimeN 24deg23prime45PrimeE c 1300 m 5vi2003 JK amp SS

16 12 3

I-Bu-10 ndash Bulgaria Stara planina Mts spring near the tourist path from Kalofer to cottage Rai under cottage Rai 42deg37prime00PrimeN 24deg58prime59PrimeE 2vi 2003 JK amp SS

ndash ndash 3

I-Bu-11 ndash Bulgaria Zap Rhodopi Mts near the Yundol village 42deg06prime17PrimeN 23deg54prime02PrimeE 1120 m 4vi2004 JK amp MK

16 ndash 3

I-Tu-12 ndash Turkey Trabzon prov Kalkanli Da lari Mts near tunnel Zigana Dag 40deg40prime36PrimeN 39deg25prime56PrimeE 1580 m 26v2004 JK amp MK

ndash ndash 3

I-Tu-13 ndash Turkey Giresum prov Giresum Da lari Mts Guumlduumll 40deg38prime43PrimeN 38deg27prime21PrimeE 880 m 29v2004 JK amp MK

ndash ndash 3

I-Ge-14 ndash Georgia Lesser Caucasus Mts Bakuriani in the forest near the WWF Education Centre 41deg45prime19PrimeN 43deg31prime15PrimeE 2vii2001 KM

ndash 21 5

I-Ge-15 ndash Georgia Greater Caucasus Mts Khevi prov N of Kazbegi Dariali canyon Gveleti along the Tergi (Terek) river near the road Tbilisi-Vladikavkaz 42deg42prime21PrimeN 44deg37prime48PrimeE 1470 m 7vii2001 KM

ndash 10 4

I-Ru-16 ndash Russia Karachay-Cherkessia Greater Caucasus Mts Dambai near first station of cable car 43deg16prime43PrimeN 41deg38prime09PrimeE 1750 m 2viii 2004 JK

ndash ndash 3

I-Ru-17 ndash Russia Karachay-Cherkessia Greater Caucasus Mts Teberda village Teberdinskii zapovednik near Teberda river 43deg26prime17PrimeN 41deg44prime08PrimeE 1320 m 3viii2004 JK amp VGO

16 ndash 3

I-Ro-18 ndash Romania Bucegi Mts near the lake Lacul Bolboci 45deg21prime15PrimeN 25deg25prime15PrimeE 1460 m 9vii2003 JK MK amp MO

ndash 22 ndash

I-Ro-19 ndash Romania Bucegi Mts Cheile Zanoagei Mari canyon 45deg19prime42PrimeN 25deg25prime16PrimeE 1320 m 9vii2003 JK MK amp MO

ndash 6 3

I-Ro-20 ndash Romania Bucegi Mts Busteni Gura Dihamului valley of Valea Seaca stream 45deg26prime35PrimeN 25deg31prime09PrimeE 990 m 9vii2003 JK

ndash 8 ndash

I-Ro-21 ndash Romania Cozia Mts 45deg18prime10PrimeN 24deg20prime12PrimeE 720 m 11vii2003 JK amp MK ndash ndash 3I-Ro-22 ndash Romania Bistri a Arcalia 47deg04prime36PrimeN 24deg21prime00PrimeE 21v2003 MK ndash ndash 2I-Ro-23 ndash Romania Turda Cheile Thurzii 46deg33prime45PrimeN 23deg40prime45PrimeE 7vii2003 JK amp MK ndash ndash 1I-Cr-24 ndash Croatia Velebit Mts Ottarje 44deg31prime45PrimeN 15deg11prime16PrimeE 960 m 18vii2003

JK amp MK ndash ndash 3

g

g

t

172 J KUBERA ET AL

copy 2006 The Linnean Society of London Botanical Journal of the Linnean Society 2006 152 169ndash195

I-Se-25 ndash Serbia Vrhpolje 44deg16prime05PrimeN 19deg26prime05PrimeE 180 m 22v2004 JK amp MK ndash ndash 3I-It-26 ndash Italy Umbria prov Terni Marmore Waterfalls 42deg32prime37PrimeN 12deg42prime58PrimeE

7v2003 JK amp MKndash ndash 5

I-It-27 ndash Italy Piemont prov W of Torino San Francesco su Avigliana 45deg4prime22PrimeN 7deg21prime374PrimeE 620 m 14v 2002 JL amp KM

ndash ndash 2

I-It-28 ndash Italy Trentino prov Val di Breguzzo 46deg02prime50PrimeN 10deg37prime25PrimeE 8vi2003 JL KM amp MS

ndash ndash 3

I-Sw-29 ndash Switzerland val di Poschiavo NE of Brusio 46deg15903N 10deg07505E 910 m 9vi2003 JL KM amp MS

ndash ndash 3

I-Sp-30 ndash Spain Leacuterida prov central Pyrenees Val drsquoAran W of Vielha Sarrat de Sepeguilha along the road from Casau to Refugi Eth Santet 42deg42prime00PrimeN 0deg46prime30PrimeE 1580 m 22vi 2002 JL

ndash ndash 2

I-Sk-31 ndash Slovakia Belianske Tatry Mts near Belianska jaskyoa cave 49deg13prime37PrimeN 20deg18prime55PrimeE 15viii2003 KM JL amp HKu

ndash ndash 3

I-Sk-32 ndash Slovakia Slovensky kras Mts near village Jasov 48deg40prime57PrimeN 20deg58prime24PrimeE 27v2003 MK

ndash ndash 3

I-Sk-33 ndash Slovakia Maleacute Karpaty Mts Borinka village Medeneacute Haacutemre 48deg15prime14PrimeN 17deg07prime15PrimeE 10v2004 JK

ndash ndash 3

I-Sk-34 ndash Slovakia Muraacutenska planina Mts near village of Muraacuteo 48deg43prime29PrimeN 20deg01prime10PrimeE 18vi2003 MK

ndash ndash 3

I-Ja-35 ndash Japan Nagano Pref Minami Azumi Co Azumi village Shimashima-dani 36deg11797 N 137deg46151E 740ndash820 m 18iv2002 JL KM HK amp SK

ndash ndash 1

I-Ja-36 ndash Japan Miyazaki Pref Higasi usuki-gun Shiiba-son Nakasuiryu on route along Kiunadani Valley 32deg07prime05PrimeN 131deg09prime13PrimeE 660 m 14vi2004 KM amp HKu

ndash ndash 3

South Korea Gangwon-do Inje-gun Buk-myeon Mt Myungdang 38deg10prime18PrimeN 128deg15prime40PrimeE 400 m vi2005 KM JL HKu et al

16 ndash ndash

South Korea Gangwon-do Pyeongchang-gun Yongpyeong-myeon Nodong-ri Mt Gyebang 37deg42prime19PrimeN 128deg29prime07PrimeE 840 m 14 vi2005 KM JL HKu et al

16 ndash ndash

Cardamine pectinata Pall ex DCP-Bu-1 ndash Bulgaria Tetevenska planina Mts valley of Zavodna river 42deg49prime21PrimeN

24deg22prime48PrimeE c 850 m 5vi2003 JK amp SS16 30 3

P-Bu-2 ndash Bulgaria Tetevenska planina Mts Tsarishina nature reserve near the cottage Vezhen 42deg46prime29PrimeN 24deg23prime08PrimeE 1560 m 5vi2003 JK amp SS

16 30 5

P-Bu-3 ndash Bulgaria Pirin Mts valley of Demyanitsa river near waterfall Yulenski skok 41deg46prime42PrimeN 23deg27prime44PrimeE 1550 m 7vi2003 JK amp SS

16 34 5

P-Bu-4 ndash Bulgaria Pirin Mts near cottage Banderitsa 41deg46prime07PrimeN 23deg25prime36PrimeE 1810 m 9vi2003 JK amp SS

ndash 30 3

P-Bu-5 ndash Bulgaria Zap Rhodopi Mts Batashka planina Batak 41deg54prime20PrimeN 24deg10prime29PrimeE 1460 m 3vi2004 JK amp MK

ndash 30 1

P-Bu-6 ndash Bulgaria Zap Rhodopi Mts Yundol 42deg06prime17PrimeN 23deg54prime02PrimeE 1120 m 4vi2004 JK amp MK

16 13 2

P-Tu-7 ndash Turkey Trabzon prov Kalkanli Da lari Mts near tunnel Zigana Dag 40deg40prime36PrimeN 39deg25prime56PrimeE 1580 m 26v2004 JK amp MK

ndash 25 5

P-Tu-8 ndash Turkey Rize prov Dog Karadeniz Da lari Mts Camlik 40deg42prime36PrimeN 40deg37prime55PrimeE 1280 m 28v2004 JK amp MK

16 28 4

P-Tu-9 ndash Turkey Rize prov Dog Karadeniz Da lari Mts IkizderendashCimil 40deg46prime08PrimeN 40deg34prime08PrimeE 510 m 28v2004 JK amp MK

16 29 5

P-Tu-10 ndash Turkey Giresum prov Giresum Da lari Mts Kuumlmbet 40deg33prime37PrimeN 38deg24prime32PrimeE 1510 m 29v2004 JK amp MK

ndash 29 3

P-Tu-11 ndash Turkey Artvin prov Balikli Da lari Mts HopandashScuruca Geccedil 41deg23prime46PrimeN 41deg31prime55PrimeN 690 m 30v2004 JK amp MK

ndash 7 5

P-Tu-12 ndash Turkey Kastamonu prov Ilgaz Da lari Mts 41deg04prime09PrimeN 33deg44prime53PrimeE 1860 m 2vi2004 JK amp MK

16 ndash 4

P-Ge-13 ndash Georgia Lesser Caucasus Mts Bakuriani in the forest near the WWF Education Centre 41deg45prime19PrimeN 43deg31prime15PrimeE 2vii2001 KM

ndash 18 7

Population code origin and collection data 2nMorphno ind

AFLP no ind

g

g

g

g

g

g

Table 1 Continued

TAXONOMY OF C IMPATIENS AND C PECTINATA 173


Figure 1 Distribution map of Cardamine impatiens (A) and C pectinata (B) showing our collection sites (circles withdots) and herbarium records (empty circles 165 records of C impatiens and 163 records of C pectinata from 15 herbaria)Total distribution range of C impatiens is indicated by dashed lines and extends further to Asia herbarium records shownare only from the area of sympatry with C pectinata Herbarium records of C pectinata marked on the map representthe whole distribution range of this taxon

A

B

174 J KUBERA ET AL


SAV SO SOM W WU (herbarium acronyms followHolmgren Holmgren amp Barnett 1990)

KARYOLOGICAL ANALYSES

Chromosome numbers were determined frommitotic metaphases of cells from root tips takenfrom germinating seeds The root tips were pre-treated with 0002 M hydroxyquinoline for 5 h Theywere fixed in a freshly prepared mixture of concen-trated ethanol and acetic acid (1 1) and hydro-lysed in a mixture of concentrated hydrochloricacid and ethanol (1 1) for 1ndash2 min Squashes weremade in a drop of 45 acetic acid under a cello-phane square (Muriacuten 1960) and stained in a 10solution of Giemsa stock solution in Soumlrensen phos-phate buffer for 1 h

MORPHOMETRIC ANALYSES

Nine quantitative vegetative characters and five floralcharacters were measured on the collected plants andthree ratio characters were derived (see Table 2) Thecharacters included in the morphometric analysis cor-responded to those usually used for the delimitation ofC pectinata and its discrimination from C impatiens(Stoyanov amp Stefanov 1948 Cullen 1965 Khatri1988 Jones amp Akeroyd 1993) as well as others thatappeared to be variable in the populations sampled inthe field Floral characters were measured on one ran-domly selected flower per plant Fresh floral parts

were attached to adhesive tape dried and scanned ona Microtek ScanMaker 9800XL Maximum values foreach floral character were measured and recorded inthe computer program CARNOY (Schols et al 2002)

We performed both hierarchical and nonhier-archical multivariate morphometric evaluation of themeasured data Hierarchical UPGMA (unweightedpair-group method using arithmetic averages) (Ever-itt 1986) cluster analysis was based on populationscharacterized by average values of the measuredcharacters The characters in the primary matrixwere standardized by zero mean and unit standarddeviation and a Euclidean coefficient was used tocompute the secondary distance matrix Principalcomponent analysis (PCA) (Sneath amp Sokal 1973Krzanowski 1990) based on individual plants andthe correlation matrix between the characters wasperformed to reduce the multidimensionality of theoriginal character space and to display the variationpattern along the first three components extractingmost of the variation To test the results from thecluster analysis based on populations canonicaldiscriminant analysis (CDA) and classificatory dis-criminant analysis (nonparametric classificatorydiscriminant analysis) (Klecka 1980) based on indi-vidual plants were computed Two groups resolvedby UPGMA (see lsquoResultsrsquo) and corresponding toC impatiens and C pectinata were defined In CDAa discriminant function was derived to maximizebetween-group variation and a histogram showingthe extent of group separation was produced To

Table 2 Eigenvectors of principal component analysis (Prin 1 Prin 2 and Prin 3) based on individuals of Cardamineimpatiens and C pectinata expressing correlation of the examined characters with the principal components (see Fig 3)total canonical structure (Can 1 correlation coefficients of the characters and canonical axis) of canonical discriminantanalysis based on individuals of C impatiens and C pectinata (see Fig 4)

Morphological character Prin 1 Prin 2 Prin 3 Can 1

Length of sepals (mm) 0728 0255 minus0146 0668Width of sepals (mm) 0538 0211 minus0165 0435Length of petals (mm) 0911 0016 minus0151 0900Width of petals (mm) 0904 0118 minus0063 0876Length of filaments (mm) 0736 minus0038 minus0321 0674Height of stem from base to the lowest peduncle of flower or fruit (cm) minus0732 0365 minus0305 minus0713Number of stems (except the main stem) 0240 0085 minus0222 0234Number of lateral inflorescences minus0269 0385 minus0566 minus0189Number of stem leaves minus0864 0162 minus0310 minus0877Number of leaflets of the middle stem leaf minus0856 0103 minus0309 minus0881Length of terminal leaflet of the middle stem leaf (LTL) (cm) 0186 0892 0211 0281Width of terminal leaflet of the middle stem leaf (WTL) (cm) 0276 0871 0116 0335Ratio LTLWTL minus0418 0443 0545 minus0390Length of lowermost lateral leaflet of the middle stem leaf (LLL) (cm) minus0016 0935 minus0140 0052Width of lowermost lateral leaflet of the middle stem leaf (WLL) (cm) 0216 0887 minus0018 0265Ratio LLLWLL minus0678 0310 0391 minus0666Ratio LTLLLL 0320 minus0008 0609 0363



reveal the correlations of characters with the canoni-cal axis the total canonical structure was com-puted The classificatory discriminant analysis wasused to assess the percentage of plants correctlyassigned to the predetermined groups The cross-val-idation procedure was used in which the classifica-tion criterion is based on N minus 1 individuals and thenapplied for the individual left out

In addition mean values standard deviations andpercentiles were computed for all characters (explor-atory data analysis) Cluster analysis was performedusing SYN-TAX 2000 (Podani 2001) For other anal-yses the SAS statistical package (SAS Institute 2000)was used

AFLP FINGERPRINTING

Genomic DNA was extracted from silica gel-driedleaves using a modified cetyltrimethylammonium bro-mide (CTAB) procedure (Doyle amp Doyle 1987) accord-ing to Schoumlnswetter et al (2002) The quality of theisolated genomic DNA was checked on agarose gelsand the amount of DNA was estimated photometri-cally (UV-160A Shimadzu) The AFLP procedure (Voset al 1995) followed the general protocol (Applied Bio-systems 1996) with minor modifications (see Schoumln-swetter et al 2004) The genomic DNA was restrictedwith MseI and EcoRI restriction endonucleases andtwo double-stranded adaptors were ligated at 37 degCfor 2 h The product was diluted in TE01 buffer(10 mM Tris 01 mM EDTA) Amplifications wereperformed in two steps using a GeneAmp PCR System9700 thermal cycler (Applied Biosystems) Preselec-tive amplification using two primers with 1-base pair(bp) extensions at the 3prime end was performed with thefollowing cycle profile initial hold at 72 degC for 2 min20 cycles at 94 degC for 1 s 56 degC for 30 s and 72 degC for2 min last hold at 60 degC for 30 min and cooling to 4 degCThe products of preselective amplifications werechecked on agarose gels and diluted in TE01 bufferThree primer combinations giving clear and reproduc-ible bands applied in previous Cardamine studies(see for example Lihovaacute et al 2004b) were used forselective amplifications EcoRI-AAG-(HEX) EcoRI-ATC-(6-FAM) EcoRI-AGC-(NED) MseI-CTG MseI-CAG and MseI-CTG (VBC Genomics Vienna Aus-tria) The polymerase chain reaction (PCR) cycle pro-file was as follows initial cycle at 94 degC for 2 min65 degC for 30 s and 72 degC for 2 min eight cycles at94 degC for 1 s 64 degC for 30 s (decreasing by 1 degC in eachcycle from 64 degC to 57 degC) and 72 degC for 2 min 23cycles at 94 degC for 1 s 56 degC for 30 s and 72 degC for2 min and the last hold at 60 degC for 30 min and cool-ing to 4 degC The AFLP fragments were loaded onto45 polyacrylamide gels with an internal size stan-dard GeneScan 500 ROX (Applied Biosystems) and

electrophoresed on an automated sequencer (ABI377) Raw data were analysed by GeneScan (AppliedBiosystems) and then by GenoGrapher program (ver-sion 160 copyMontana State University 1999 httphordeummsumontanaedugenographer) The pres-ence or absence of AFLP fragments was scored foreach sample (only well-scorable and unambiguousfragments were analysed) and transferred into abinary matrix

AFLP DATA ANALYSES

Genetic variation was analysed from both a taxonomicand phylogeographical perspective

1 First principal coordinate analysis (PCoA) andneighbour-joining (NJ) analysis were performedon the entire dataset of 147 samples to obtain thegeneral view on the AFLP variation pattern partic-ularly to address the genetic differentiation betweenpopulations morphologically assigned to C impatiensand C pectinata For PCoA the secondary matrixwas computed using Jaccardrsquos coefficient and theanalysis was performed by SYN-TAX 2000 (Podani2001) NJ analysis was carried out using Nei amp Lirsquos(1979) genetic distance in the PAUP program (ver-sion 40b10 Swofford 2003) with the bootstrapoption of 5000 replicates Three accessions ofC lazica Boiss were used to root the tree

2 To document the genetic divergence between thetwo taxa we additionally calculated the totalnumber of fragments per taxon the number of poly-morphic fragments the number of private fixedfragments (those restricted to the taxon andpresent in all of its individuals) and the number ofprivate fragments (those restricted to the taxon butnot necessarily present in all of its individuals)

3 The geographical structure of genetic variationwas explored by analyses of molecular variance(AMOVAs) using ARLEQUIN (version 2000Schneider Roessli amp Excoffier 2000) The originalbinary matrix was divided into two matrices ofC impatiens and C pectinata respectively and theAMOVAs were run separately for each of these twotaxa Euclidean distances were computed betweenall pairs of AFLP phenotypes to produce distancematrices Total genetic variation was partitionedinto the following hierarchical levels between indi-viduals within populations between populationsand between regions Different groupings of popu-lations were tested following those suggested byPCoA and NJ analysis or by phytogeographicalareas One thousand permutations were run toobtain test statistics

4 Finally to estimate the genetic diversity withinpopulations and geographical regions the followingdiversity measures were calculated using POP-

176 J KUBERA ET AL


Figure 2 Cluster analysis (unweighted pair-group method using arithmetic averages UPGMA) of populations of Car-damine impatiens (I) and C pectinata (P) based on 17 morphological characters

Dis

sim

ilarit

y

6

5

4

3

2

1

0

I-Bu-

1

I-Bu-

3

I-Bu-

4

I-Ro-

20

I-Bu-

2

I-Ro-

18

I-Ge-

15

I-Bu-

5

I-Bu-

8

I-Ro-

19

I-Bu-

7

I-Ge-

14

I-Bu-

6

I-Bu-

9

P-B

u-1

P-B

u-2

P-B

u-4

P-B

u-6

P-B

u-5

P-B

u-3

P-G

e-13

P-T

u-10

P-T

u-11

P-T

u-9

P-T

u-8

P-T

u-7

C impatiens

C pectinata

GENE 132 (Yeh et al 1997) the total number ofAFLP fragments in a populationregion (fpop) themean number of AFLP fragments per individual ina population (find) the number and percentage ofpolymorphic fragments (Ppop) the number of rare(fr fragments present in less than 10 of the inves-tigated individuals belonging to the taxon) private(fpr) and private fixed (fpr-f) fragments and Shan-nonrsquos diversity index HSh = ndashΣ(pi ln pi) where pi isthe relative frequency of the ith fragment (Leg-endre amp Legendre 1998) As the numbers of indi-viduals per population analysed for AFLPs werenot equal but ranged from one to nine individualsthese diversity measures were calculated onreduced datasets that included three (the mostcommon number) or exceptionally two individuals(in cases in which only two individuals were sam-pled) per population From populations in whichmore individuals were sampled three individualswere randomly selected

RESULTS

CHROMOSOME NUMBERS

Chromosome numbers were counted in eight popula-tions of C impatiens and seven populations ofC pectinata The diploid chromosome number 2n = 16was determined in all cases (Table 1)


UPGMA cluster analysis based on 17 morphologicalcharacters resulted in a dendrogram with two mainclusters corresponding to populations traditionallyunderstood as C impatiens and C pectinata respec-tively (Fig 2) Further clustering in C pectinatareflected the geographical origin of the analysedpopulations (Balkan and TurkeyCaucasus region)whereas clustering within C impatiens did not showany pattern The ordination diagram of PCA based onindividual plants (Fig 3) showed two compact group-ings separated along the first axis in accordance withthe results of cluster analyses Only a minute overlaprepresented by few plants was present The followingcharacters were strongly correlated with the firstcomponent axis (in descending order) length of petalswidth of petals number of stem leaves number of leaf-lets of the middle stem leaf length of filaments heightof stem and length of sepals (Table 2) No furthergroupings were apparent along the second or thirdaxes although there was some variation within bothgroups especially in C impatiens

The histogram of CDA (Fig 4) based on individualplants and two groups as defined by cluster analysisshowed a clear separation between the populationscorresponding to C impatiens and C pectinata Thecharacters highly correlated with the canonical axiswere the same as revealed in PCA (Table 2) It is



apparent that there are no (or only very few) plantindividuals that can be considered as morphologicallyintermediate between C impatiens and C pectinataAccordingly in the classificatory discriminant analy-sis percentages of correctly classified individuals were100 without any misclassified individuals

The results of exploratory data analysis ofC impatiens and C pectinata (Fig 5) showed thatthese two taxa differed in several characters with anoverlap only in extreme values of a few individualsAgain the length of petals width of petals number ofstem leaves number of leaflets of the middle stem leaflength of filaments height of stem length of sepalsand the ratio LLLWLL (length of lowermost lateralleaflet of the middle stem leafwidth of lowermost lat-eral leaflet of the middle stem leaf) were the charac-ters best differentiating between these two taxa

AFLP ANALYSES

PCoA based on AFLP data resulted in two groupingsplaced on opposite sides of the first axis (figurenot shown) corresponding to C impatiens andC pectinata respectively The pronounced geneticdifferentiation was also apparent from the NJtree where two main clusters of C impatiens andC pectinata with high bootstrap values were resolved(Fig 6) Of the 214 fragments scored in 147 individu-als (excluding C lazica used to root the tree) 198

(93) were polymorphic In C impatiens 153 frag-ments were resolved 130 (85) were polymorphic 88were private and seven were private fixed InC pectinata 119 fragments were found 95 (80) werepolymorphic 53 were private and eight were privatefixed

Apart from the clear taxonomic structure resolvingC impatiens and C pectinata as two genetically dif-ferentiated entities the NJ tree suggested furthergeographical patterns within both taxa Two mainclusters in C pectinata corresponded to populationsfrom (1) the Balkan Peninsula and western and cen-tral Turkey and (2) eastern Turkey and the Caucasushowever the clusters received very low bootstrap sup-port Within both clusters further clustering againreflected the geographical origin of the accessionsSimilarly three main clusters were apparent withinC impatiens (1) Japan (2) eastern group of Turkeyand Caucasus and (3) western group of the BalkanPeninsula and other parts of Europe (Fig 6)

Several three-level AMOVAs were performed toassess partitioning of the overall genetic variation andto test the suggested geographical structure withinAFLP data (Table 3) Two to five geographical group-ings were suggested for both taxa to compare thebetween-region variance In C impatiens about 20ndash26 of the total variation was accounted for by varia-tion within populations and 30ndash38 by variationbetween populations By partitioning populations into

Figure 3 Principal component analysis of individuals of Cardamine impatiens (cubes) and C pectinata (pyramids) basedon 17 morphological characters (see Table 2) The first three axes explain 3557 2329 and 1018 of the total variationrespectively

-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL


two main geographical groups the western and east-ern groups (Japanese accessions were excluded fromAMOVA) almost 50 of the overall genetic variationwas attributed to the differentiation between thesetwo areas By partitioning the populations intosmaller groups the geographical resolution wasalways weaker (Table 3) In C pectinata the highestbetween-region differentiation was achieved whenconsidering three geographical groups (Balkan Penin-sula central Turkey eastern Turkey together with theCaucasus) representing almost 61 of the total vari-ation This value decreased slightly (to 58) when twogroups were considered eastern Turkey together withthe Caucasus as one group and the remaining popu-lations as the second group (Table 3)

Table 4 presents genetic diversity estimates for theanalysed populations and geographical regions whichare also shown in Figure 7 In most cases the numberof individuals analysed was the same as the number ofdetected AFLP phenotypes Only in a single popula-tion of C impatiens (I-Ge-15 Georgia Gveleti) did allthree individuals share the same AFLP phenotype InC pectinata two pairs of individuals sharing the sameAFLP phenotype were found In C impatiens several

populations with high levels of genetic diversity(reflected by the high percentage of polymorphic frag-ments number of rare and private fragments andhigher HSh values) were identified which were concen-trated to the western part of the studied area (westernCarpathians regions south of the Alps) A few popu-lations with higher genetic variation or with severalprivate AFLP fragments were also found scattered inBulgaria Romania and the Caucasus In C pectinatathe eastern populations (located along the Black Seacoast in northern Turkey and the Caucasus) displayedhigh genetic diversity with several private fragmentsin contrast with Bulgarian populations with reducedgenetic variation In populations of the latter regionhowever a few private fragments were also present

DISCUSSION

TAXONOMIC POSITION OF C PECTINATA AND ITS MORPHOLOGICAL AND MOLECULAR DIFFERENTIATION

Different taxonomic treatments can be found in theliterature with regard to the recognition and taxo-nomic status of the taxon originally described as

Figure 4 Canonical discriminant analysis of individuals of Cardamine impatiens (white) and C pectinata (cross-hatched)based on 17 morphological characters (see Table 2)

FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66



Figure 5 Variation of selected morphological characters of Cardamine impatiens and C pectinata Rectangles define 25thand 75th percentiles horizontal lines show median whiskers are from 10th to 90th percentiles and asterisks show extremevalues LLLWLL length of lowermost lateral leaflet of the middle stem leafwidth of lowermost lateral leaflet of the middlestem leaf

0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

C pectinata (Assenov 1970 Khatri 1988 Kit Tan2002) Several authors did not accept it as a separatespecies but placed it within C impatiens (Stoyanov ampStefanov 1948 Zohary et al 1980 Jones amp Akeroyd1993) Although morphological differences betweenthese two taxa have been reported the existence ofmorphological intermediates has also been assumed(Cullen 1965 Kit Tan 2002) According to Flora Euro-paea (Jones amp Akeroyd 1993) and national floras fromthe relevant areas (eg Stoyanov amp Stefanov 1948Grossgeim 1950 Cullen 1965 Assenov 1970 KitTan 2002) several morphological characters such asthe height of plants number of leaflets of the stemleaves shape of the leaflets length of the petalsbranching of the stem width of the siliquae length ofthe style and the position of the siliquae in the inflo-rescence can distinguish between C impatiens andC pectinata (often treated as a subspecies or varietywithin the former) Until our study however nodetailed morphological investigation has been per-formed Our results based on a large amount of plantmaterial showed two morphologically clearly differen-tiated entities corresponding to C impatiens andC pectinata We have not recorded any morphologi-cally intermediate or uncertain populations whichcannot be classified reliably This was confirmed at thelevel of individual plants and also from localities inwhich both taxa grew in close proximity Morphologi-cal characters which were resolved in our analyses asdiagnostic mostly corresponded to those reported by

the authors of several taxonomic treatments (forreferences see above) Fruit characters were notincluded in our detailed morphometric evaluation asthe population samples were collected at floweringtime Nevertheless we studied numerous fruitingspecimens in herbaria C pectinata typically pos-sesses secund siliquae and the resulting fruitinginflorescence shape can be considered as a characterdistinguishing it from C impatiens however excep-tions occur In addition to the traditionally reporteddifferentiating characters our study showed differ-ences in the length of the sepals width of the petalsand length of the filaments The extreme values of thediscriminating characters may overlap in some cases(see Fig 5) but by using a combination of a few char-acters it is always possible to identify C impatiens orC pectinata without doubt

In populations of both C impatiens andC pectinata plants with hairy siliquae are seldomfound In the past they were classified either as a sep-arate species C dasycarpa M Bieb or as varieties ofeither C impatiens (var eriocarpa DC) or C pectinata(var trichocarpa N Busch) Following the lectotypifi-cation in the present paper C dasycarpa is a synonymof C impatiens and two other names fall into the syn-onymy of C pectinata The character lsquohairness of sili-quaersquo apparently has no taxonomic value In additionapetalous plants very often occur in populations ofC impatiens They were named by Gilibert (1782) asC apetala however this name is currently treated as

180 J KUBERA ET AL


Figure 6 Neighbour-joining tree of amplified fragment length polymorphism (AFLP) data including 95 accessions ofCardamine impatiens and 52 accessions of C pectinata For explanation of the accession labels see Table 1 Bootstrapvalues are shown above the branches Cardamine lazica was used to root the tree

I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes



Table 3 Analysis of molecular variance (AMOVA) of amplified fragment length polymorphism (AFLP) data performedwith different geographical groupings within both Cardamine impatiens (AndashD) and C pectinata (EndashH) C Partitioning ofpopulations in the Caucasus and eastern Turkey follows the pattern resolved by the neighbour-joining tree E TurkeyCaucasus (1) represents populations I-Tu-13 I-Ru-16 -17 E Turkey Caucasus (2) represents populations I-Tu-12 I-Ge-14 -15 Values discussed in the text are marked in bold

Grouping Source of variation dfSum ofsquares

Variancecomponents

of totalvariance

Cardamine impatiensA [Turkey Caucasus][Europe] Between groups 1 24388 705 4932dagger

Between populations 27 43625 443 3099daggerWithin populations 59 16608 281 1969dagger

B[TurkeyCaucasus][Carpathians][Balkan] Between groups 3 30653 403 3702dagger[western Europe] Between populations 25 37360 404 3711dagger

Within populations 59 16608 281 2587daggerC [E Turkey Caucasus (1)][E Turkey Caucasus (2)] Between groups 4 33928 416 3855dagger[Carpathians][Balkan][western Europe] Between populations 24 34085 382 3537dagger

Within populations 59 16608 281 2608daggerD [Caucasus][Turkey][Carpathians][Balkan] Between groups 4 32243 390 3628dagger[western Europe] Between populations 24 35769 404 3755dagger

Within populations 59 16608 281 2617daggerCardamine pectinataE [Turkey Caucasus][Bulgaria] Between groups 1 16799 648 5109


F [E Turkey Caucasus][central Turkey][Bulgaria] Between groups 2 23422 762 6092daggerBetween populations 9 9837 188 1506daggerWithin populations 39 11718 300 2402dagger

G [E Turkey Caucasus][central TurkeyBulgaria] Between groups 1 20225 738 5772Between populations 10 13033 240 1879daggerWithin populations 39 11718 300 2349dagger

H [Caucasus][Turkey][Bulgaria] Between groups 2 19462 509 4538Between populations 9 13796 312 2783daggerWithin populations 39 11718 300 2679dagger

df degrees of freedomP lt 001daggerP lt 0001

invalid (see lsquoTaxonomic conspectusrsquo) Moench (1794)apparently intended to publish the name C apetalaagain for apetalous plants of C impatiens but theonly original element connected with this name anillustration in Flora Danica (Muumlller 1777 table 735)depicts undoubtedly C flexuosa With This variationhas no taxonomic value

From our AFLP data C impatiens and C pectinataare also genetically well differentiated The largenumber of private AFLP fragments as well as theoverall variation pattern illustrated by PCoA and theNJ tree show pronounced genetic differentiationbetween these two taxa These results are in clearaccordance with morphometric data and provide con-vincing evidence that C impatiens and C pectinatashould be treated as two different species We did notfind any signs of interspecific gene flow no intermedi-

ate AFLP phenotypes were seen on the ordination dia-gram of PCoA and no morphological intermediateswere observed Our recent study on phylogenetic rela-tionships between European Cardamine diploid taxausing DNA sequence data from three different mark-ers (ITS nrDNA trnL-trnF of cpDNA and nuclearsingle-copy CHS gene) also included a few accessionsof C impatiens and C pectinata They were resolvedas two closely related sister taxa forming a well-supported and distinct lineage amongst the otheranalysed taxa (Lihovaacute Shimizu amp Marhold 2006)

Thirty original chromosome number records of2n = 2x = 16 have been published for C impatiensacross its distribution range (for details see Karyolog-ical database of the genus Cardamine http147213100117cardamineindexphp Kudera Valkoamp Marhold 2005) This is in agreement with our

182 J KUBERA ET AL


Table 4 Distribution of amplified fragment length polymorphism (AFLP) fragments and genetic diversity measures forthe populations and regions analysed (see also Fig 7) The population codes follow Table 1

Region Pop N fpop find Ppop fr fpr fpr-f HSh

Cardamine impatiensTurkey 87 25 (1656) 0104

I-Tu-12 3 76 73 7 (265) 3 ndash ndash 0030I-Tu-13 3 76 74 5 (331) 8 4 4 0021

Caucasus 92 34 (2252) 0113I-Ge-14 3 75 72 4 (265) 3 ndash ndash 0017I-Ge-15 3 74 74 0 3 1 1 0I-Ru-16 3 75 72 7 (464) 3 ndash ndash 0030I-Ru-17 3 82 74 18 (1192) 6 1 ndash 0076

Balkan 96 44 (2914) 0115I-Bu-2 3 74 68 10 (662) 3 ndash ndash 0042I-Bu-3 3 72 70 6 (397) 1 1 ndash 0025I-Bu-4 3 72 67 8 (53) 2 1 ndash 0034I-Bu-7 3 72 69 5 (331) 1 ndash ndash 0021I-Bu-8 3 70 67 7 (464) 2 1 ndash 0029I-Bu-9 3 76 73 5 (331) 3 ndash ndash 0021I-Bu-10 3 70 68 3 (199) 0 ndash ndash 0013I-Bu-6 2 72 71 3 (199) 2 ndash ndash 0014I-Bu-11 3 68 67 2 (132) 0 ndash ndash 0008I-Cr-24 3 69 67 4 (265) 2 ndash ndash 0017I-Se-25 3 73 69 9 (596) 2 1 1 0038

E S Carpathians 82 27 (1788) 0098I-Ro-19 3 71 67 9 (596) 2 ndash ndash 0038I-Ro-21 3 74 69 10 (662) 2 ndash ndash 0042I-Ro-22 2 72 71 3 (199) 2 2 ndash 0014

W Carpathians 97 42 (2781) 0133I-Sk-31 3 88 75 23 (1523) 9 4 ndash 0097I-Sk-32 3 68 66 5 (331) 2 1 ndash 0021I-Sk-33 3 75 69 11 (728) 2 ndash ndash 0046I-Sk-34 3 77 70 13 (861) 2 ndash ndash 0055

W Europe 105 37 (2450) 0125I-Sw-29 3 82 75 13 (861) 4 3 2 0055I-It-28 3 77 68 17 (1126) 4 ndash ndash 0072I-It-27 2 76 72 8 (53) 4 ndash ndash 0037I-It-26 3 72 71 2 (132) 2 ndash ndash 0008I-Sp-30 2 73 70 6 (397) 3 ndash ndash 0028

Cardamine pectinataCaucasus amp 107 67 (5726) 0216Turkey P-Ge-13 3 67 62 10 (855) 4 3 ndash 0054

P-Tu-11 3 65 62 7 (598) 3 2 ndash 0037P-Tu-8 3 64 63 3 (256) 1 ndash ndash 0016P-Tu-9 3 70 66 8 (684) 2 ndash ndash 0043P-Tu-7 3 78 65 24 (2051) 9 5 ndash 0128P-Tu-10 3 71 63 16 (1368) 7 6 ndash 0086P-Tu-12 3 77 70 12 (1026) 6 5 2 0064

Bulgaria 79 16 (1368) 0050P-Bu-1 3 73 69 8 (684) 1 1 ndash 0043P-Bu-2 3 70 68 4 (342) 0 ndash ndash 0021P-Bu-3 3 71 68 5 (427) 2 1 ndash 0027P-Bu-4 3 74 69 9 (769) 5 4 ndash 0048P-Bu-6 3 69 68 2 (171) 1 ndash ndash 0012

Pop population code N number of investigated individuals (reduced dataset with two to three individuals per populationsee lsquoMaterial and methodsrsquo) fpop total number of fragments in a populationregion find mean number of fragments perindividual in a given population Ppop number and percentage of polymorphic fragments in a populationregion fr numberof rare fragments (present in less than 10 of individuals) fpr number of private fragments fpr-f number of private fixedfragments HSh Shannonrsquos diversity index for a populationregion Three classes of HSh values were established to discussthe diversity patterns high values are in bold medium values are in bold italic and low values are in normal type



results the only reliably documented chromosomenumber for this species Three authors have also pub-lished the tetraploid level for C impatiens The tetra-ploid record by Hindaacutekovaacute (in Maacutejovsky et al 1974)published from Slovakia refers in fact to C flexuosa(based on the revision of the voucher specimen Mar-hold 1994b) We have not been able to trace thevoucher specimens of the tetraploid chromosome num-bers published by Gohil Ashruf amp Raina (1981) andJee Dhar amp Kachroo (1983) from the area of KashmirTherefore we cannot consider these data as reliableThese records may well refer to another taxon Inthe account of the Cruciferae of Continental NorthAmerica (Rollins 1993) the tetraploid number wasreported but without the indication of any particularlocality and thus this record cannot be considered as

reliable For C pectinata the only chromosome num-ber record was published from Bulgaria (Anchev ampGoranova 1999) which is in accordance with ourresults confirming the diploid level from seven popu-lations (Table 1)

GEOGRAPHICAL DISTRIBUTION AND ECOLOGY OF C PECTINATA

Based on the revision of herbarium specimens depos-ited in relevant herbaria we present the distributionarea of C pectinata (Fig 1 Appendix) It covers moun-tain ranges in the Balkan Peninsula (BulgariaGreece) northern Turkey the Caucasus region andnorthern Iran There is one herbarium record originat-ing from a locality in southern Turkey near the border

Figure 7 Sample sites of Cardamine impatiens (A) and C pectinata (B) included in our amplified fragment lengthpolymorphism (AFLP) study Population codes follow Table 1 Intrapopulational genetic diversity as assessed by Shannonrsquosdiversity index (HSh) is indicated by different circle sizes (small circle low HSh medium circle medium HSh large circlehigh HSh see Table 4) The number of private AFLP fragments is shown by lsquo+rsquo Arrows delimit geographical groups thatshow the most pronounced genetic differentiation (analysis of molecular variance AMOVA see Table 3 values in bold) InC pectinata two almost equal partitionings are indicated

++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4

+ central groupwestern group eastern groupeastern group

western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh

+ number ofprivate fragments

I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+

western group eastern group

184 J KUBERA ET AL


with Syria which is distant from any other reportsfrom northern Turkey (see Fig 1 Appendix) The spec-imen was collected in 1913 by M Haradjian in theAmanus Mountains (Nur Da lari) that belonged atthat time to Syria nowadays to the province of Hatayin Turkey It was identified by Rechinger asC impatiens but morphologically it undoubtedly cor-responds to C pectinata As a result of the large geo-graphical disjunction and only a single record fromthat area we may consider this record as doubtfulHowever M Haradjian is known by his collectionsfrom Syria and Cyprus (Lanjouw amp Stafleu 1957) andwe are not aware of any of his collections from theknown distribution area of C pectinata Therefore itseems that the localization of the origin of this speci-men is correct and extends the known distributionarea of C pectinata to the area of southern Turkey

From the consultation of numerous floras from therelevant area a similar ecology has been reported forC impatiens and C pectinata Both species usuallygrow on moist sites in shady forests however differ-ences reported between C impatiens and C pectinatain terms of altitudinal range and forest types are con-troversial Our field observations on the other handindicate a clear and consistent pattern in terms of eco-logical differentiation C pectinata occurs in conifer-ous or mixed (AbiesndashFagus) forests we have not foundany locality in an entirely deciduous forest The alti-tudinal range was recorded to be 690ndash1860 m with asingle locality at 505 m where only a few plants wereobserved apparently originating from a populationgrowing at higher elevation C impatiens on the otherhand occurs either in lowland deciduous forests or inmixed (AbiesndashFagus) forests but has not beenobserved in an entirely coniferous forest It is usuallyobserved at lower altitudes from 180 to 1750 m pop-ulations in the upper altitudinal range occur only inthe Caucasus Mountains with surrounding peaksreaching 5000 m Both species can meet in mixed for-ests and indeed we have observed three localities inwhich they are found in close vicinity (populations I-Bu-11 and P-Bu-6 I-Tu-12 and P-Tu-7 I-Ge-14 and P-Ge-13 Table 1) Even in these cases however theyshowed microspatial separation indicating differ-ences in their ecological preferences

PHYLOGEOGRAPHICAL HYPOTHESES

Different distribution areas occupied by C impatiensand C pectinata and the concentration of C pectinatapopulations in assumed glacial refugia (Hewitt 2004)led us to explore the phylogeographical patternswithin these species AMOVA and the assessment ofgenetic diversity have been applied to reveal the geo-graphical partitioning of genetic variation to identifythe genetically most diverse populations (potential

g

refugial populations serving as source populations forpostglacial colonization) and to track postglacialmigrations In C pectinata strong genetic differentia-tion was observed between the populations from east-ern TurkeyCaucasus central Turkey and Bulgaria(see Fig 7) indicating long-term isolation and restric-tion of gene flow between these three regions Popu-lations from mountain ranges along northern Turkey(P-Tu-12 -10 -7) and a population from the LesserCaucasus (P-Ge-13) displayed the highest HSh valuesaccounting for both the number of AFLP fragmentsand their frequency The same populations also har-boured most private fragments These genetic pat-terns are in clear accordance with the assumedlocation of glacial refugia along the Black Sea coast asshown by several authors (eg Hampe et al 2003Marcussen 2003 Krebs et al 2004) Survival of pop-ulations within this area and altitudinal shifts duringglacials and interglacials may have enabled them toretain considerable genetic variation and accumulateprivate fragments (see Petit et al 2003) C pectinatacurrently grows at higher altitudes within this areathus during colder periods it may have found suitablehabitats in the same or neighbouring areas at loweraltitudes Indeed palaeoclimatological and palynolog-ical data (Adams 1997) suggest that such suitablehabitats were available within this area Themountainous distribution of C pectinata also explainsthe strong geographical differentiation revealed byAMOVA Gene flow between the populations occupy-ing different mountain ranges is more restricted thanif the species was continuously distributed The threegenetically differentiated regions (Fig 7) coincide wellwith the current distribution pattern showing distri-butional gaps in the lowlands separating these regions(the area of north-west Turkey west of Istanbul andcentral Turkey Fig 1) Bulgarian populations locatedat the western distribution border of this species onthe other hand were revealed as less diverse but stillharboured private fragments not present in the popu-lations from Turkey and the Caucasus This indicatesthat the mountainous area of Bulgaria has been colo-nized postglacially but probably not from the refugiain Turkey According to the reconstruction of thevegetation composition in the Balkans during the lastglacial maximum there have been areas with a con-tinuous presence of both coniferous and deciduous treespecies (Willis 1994) A high diversity of tree specieshas been documented eg in Ioannina (north-westGreece) C pectinata may have survived the unfavour-able climatic conditions in such refugial populationsUnfortunately we have not sampled populations fromGreece that may have served as source populations forthose in more northern areas of Bulgaria To obtainmore detailed insights into the phylogeography of thisspecies more individuals per population larger sam-



pling within the analysed regions and sampling inareas not covered here (Greece northern Iran theGreater Caucasus) are needed

The distribution range of C pectinata is intriguingbecause in contrast with the widespread C impatiensit is much smaller and restricted to refugial areas andadjacent regions but still within the range of the lat-ter Considering their sister relationships we canassume two speciation scenarios from their commonancestor Allopatric speciation may have occurredpreglacially near the current distribution ofC pectinata and the current sympatry may have orig-inated as a result of their secondary contact Alterna-tively sympatric speciation may have occurred but inthis case strong genetic barriers must have been gen-erated that have prevented gene flow Detailed geneticand genomic studies may reveal the mechanisms oftheir reproductive isolation Self-pollination has beenreported for C impatiens (Kimata 1983) but assum-ing from the rather high intrapopulation variationoutcrossing should also occur

Cardamine impatiens in contrast withC pectinata exhibits only a weak phylogeographicalstructure within the studied area AMOVA revealsthe eastndashwest differentiation (see Fig 7) but less pro-nounced than in C pectinata and further genetic dif-ferentiation within these two areas is not supportedPopulations harbouring the highest genetic diversityare scattered across the investigated area They arepresent mainly in the previously glaciated areas or intheir vicinity where they may have only hardly sur-vived during the glacials (the Greater CaucasusMountains the Alps and the Carpathians) There-fore we can hypothesize that the high genetic diver-sity revealed in these regions has been generatedthrough the contact between colonization routes fromdifferent refugia as found in many other species(Petit et al 2003) Alternatively the efficient spreadand mixing between different colonization routes mayhave completely blurred any phylogeographicalpattern that may have resulted from postglacialmigrations

TAXONOMIC CONSPECTUS OF C IMPATIENS AND C PECTINATA

CARDAMINE IMPATIENS L SP PL 655 1753

Ind loc lsquoHabitat in Europae nemoribus ad radicesmontiumrsquoLectotype LINN no 8359 (Jafri 1973 169)equiv Ghinia impatiens (L) Bubani Fl Pyren 3 1601901

= C brachycarpa Opiz Naturalientausch 11 4111826

Ind loc lsquoIn monte wysokaacute hora ad Karlstein C herboeumlmiaersquoType Not designated

= C impatiens f apetala O E Schulz Bot Jahrb Syst32 455 1903Ind loc lsquoChina centralis prov Szechrsquouan prNanchrsquouan leg v Rosthorn 1891 n 2049 leg idem n2048 (H B) prov Hupeh leg A Henry 1885ndash88 n7855 (H B H C) in m Tientai leg E Faber 1889 (HB) Japonia leg Rein (H B)rsquoType Not designated

= C dasycarpa M Bieb Fl Taur-Cauc 3 437 1819Ind loc lsquoHabitat in Iberiacirc occidentalirsquoLectotype (designated here) ex herb Steveni 1809(LE)equiv C impatiens lsquoprolesrsquo dasycarpa (M Bieb) O ESchulz Bot Jahrb Syst 32 457 1903equiv C impatiens var dasycarpa (M Bieb) T Y Cheo ampR C Fang Bull Bot Lab N-E Forest Inst Harbin 621 1980

= C impatiens lsquoprolesrsquo elongata O E Schulz BotJahrb Syst 32 459 1903Ind loc lsquoAsia Affghanistan leg Griffith ante 1861 n1358 (H B H C H V) pr Otipore leg idem (HBoiss) Tibet leg Huumlgel n 185 (H V) Himalaya bor-occid 1670ndash3340 m leg T Thomson (H B H C HV) pr Abetamorsch leg Stolitzka 2000ndash2340 m (H V)Kashmir 2670ndash3000 m leg J F Duthie 1892 (H B)Garhwal in Nila Valley pr Phulaacuteldaru 3340ndash3370 mleg idem 1883 n 916 (H B Boiss) pr Sing-jari3000 m legg R Strachey et J E Winterbottom n 3pr Rogila 3370 m legg iidem n 10 Kumaon pr Dew-ali 2830 m legg iidem n 9 (H C H P Ac) Sikkim2000ndash3340 m leg J D Hooker (H B H Boiss H CH V) Mandschuria austr-litoralis ad Olga Bai leg FSchmidt 1861 (H P Ac)rsquoLectotype (designated here) Garhwaacutel Sing-jari9000 feet sa leg R Strachey et J E Winterbottomn 3 (GH)

= C impatiens var pilosa O E Schulz Bot JahrbSyst 32 459 1903Ind loc lsquoTurkestania in distr Akdishan et Naman-gan leg D Litwinow 1899 pr Wernij leg F Killomann1888 (H P Ac) Altai leg Ledebour (H P) Chinaprov Schensi sept in m Hua-tzo-pin leg Gius Giraldi1894 (H Biondi n 443 in H B) prov Szechrsquouan leg vRosthorn 1891 n 2054 (H B)rsquoLectotype (designated here) Turkestania distrAkdishan fl Kara-Lesigur 30v1899 leg D Litvinow(LE)

= C impatiens var angustifolia O E Schulz BotJahrb Syst 32 459 1903

186 J KUBERA ET AL


Ind loc lsquoCaucasus centralis ad fl Aragwa legLagowski (H P)rsquoHolotype Am Aragwa leg Dr Lagowski (LE)

= C impatiens var obtusifolia Knaf Flora 19 2941864Ind loc lsquoBei Hallstadt in OberoumlsterreichrsquoType Not designatedequiv C impatiens f obtusifolia (Knaf) Markgr in HegiIll Fl Mitteleurop ed 2 4(1) 205 1958

= C impatiens var microphylla O E Schulz BotJahrb Syst 32 460 1903Ind loc lsquoIns Sachalin leg F Schmidt 1860 (H P)rsquoHolotype Insula Sachalin zaliv Estaing 1860 FSchmidt (LE)

= C impatiens var acutifolia Knaf Flora 19 2941846Ind loc Not statedType Not designated

= C impatiens f macropetala (Boiss) O E SchulzBot Jahrb Syst 32 459 1903equiv C impatiens γ macropetala Boiss Fl Orient 311888Ind loc lsquoHab secus torrentes vallis subalpinaeDjimil Ponti Lazici 6000rsquo (Bal)rsquoHolotype Bords des torrents de la valleacutee sous-alpinede Djimil (Lazistan) vers 2000 meacutetre drsquoaltitudeJuillet-Aoucirct 1866 leg Balansa (G)

= C impatiens var minima Schur Verh Naturf Vere-ins Bruumlnn 15 77 1876Ind loc lsquoAuf buschigen Abhaumlngen am Wege nachJundorf unweit der neuen Welt bei Bruumlnn mit Linariaarvensis Juni 1871 Auch in Siebenbuumlrgen 1850rsquoType Not designated

ndash C apetala Gilib Fl Lith 1 42 1782 nom inval(opera uttique opressa)ndash C impatiens var communis N Busch Fl Sib i DalVostoka 2 243 1915 nom inval (Art 243)

Excluded name C apetala Moench Meth PlantasHorti Bot et Agri Marburgensis 259 1794Ind loc lsquoh frequens am Rhinberg prope CaldernrsquoLectotype (designated here) Muumlller Fl Dan 5131777 table 735= C flexuosa With

Description Biennial or rarely annual herb Height ofstem from base to the lowest peduncle of flower or fruit(15ndash) 18ndash75 (minus85) cm Stem erect sometimes flexuoussimple or very rarely branched glabrous Rosetteleaves shrivelled by anthesis pinnate sparsely hairywith four to eight pairs of petiolulate circular-ovate

lobate or sometimes pinnatipartite leaflets Caulineleaves (7ndash) 9ndash25 (minus28) auriculate pinnate or some-times pinnatisect above with (7ndash) 9ndash17 (minus19) denticu-late to lobed or pinnatipartite leaflets Terminalleaflets (07ndash) 09ndash28 (minus33) times (02ndash) 03ndash12 (minus15) cmpetiolulate or sessile lanceolate to ovate lanceolateLateral leaflets (05ndash) 07ndash23 (minus26) times (018ndash) 02ndash10(minus14) cm petiolulate or sessile lanceolate to ovateInflorescence racemose with 0ndash14 (minus17) lateral inflo-rescences Sepals oblong (10ndash) 12ndash23 (minus25) times (03ndash)05ndash08 (minus09) mm margin membranous petals (17ndash)19ndash35 (minus37) times (05ndash) 06ndash11 (minus13) mm white oblan-ceolate apex rounded Stamens 6 anthers greenndashyel-low filaments (15ndash) 17ndash29 (minus34) mm long stigmanot conspicuous Pedicels and siliquae patent or erect-patent

Chromosome number 2n = 2x = 16

Ecology Moist sites in shady forests lowland decidu-ous forests or mixed forests up to 2400 m (Caucasus)

Distribution area Eurasia introduced to NorthAmerica and southern Africa

C PECTINATA PALL EX DC SYST NAT 2 264 1821

Ind loc lsquoHab In Persiacirc Pallas (v s sp in h Lamb)rsquoLectotype (designated here) E Persia Herb Pallas(BM)equiv C impatiens ssp pectinata (Pall ex DC) Stoj ampStef Fl Batildelg ed 3 503 1948equiv C impatiens var pectinata (Pall ex DC) TrautvTrudy Imp S-Peterburgsk Bot Sada 4 (1) 105 1876

= C impatiens var eriocarpa DC Syst Nat 2 2621821Ind loc lsquoin Iberiacirc occidentalirsquoHolotype Imeretia parte var impatientes Iberiaem[isit] Steven 1820 (G-DC)

= C oxycarpa Boiss Ann Sci Nat (Paris) ser 2 1756 1842Ind loc lsquoN 113 Alpes Laristan 4119 ZarinkoursquoLectotype (designated here) Mt Zarinkou Aucher-Eloy-Herbier drsquoOrient no 4119 (G)

= C pectinata var trichocarpa N Busch Fl CaucasicaCritica 3 340 1904ndash10Ind loc lsquoHab in umbrosis silvaticis et ad rivulosregionis subalpinae Caucasi occidentalis et Transcau-casiae occidentalis Exsicc Aucher-Eloy HerbierdrsquoOrient no 4119 B Balansa Plantes drsquoOrient(1866) no 1361rsquo



Lectotype (designated here) Mt Zarinkou Aucher-Eloy-Herbier drsquoOrient no 4119 (G)

= C lucorum Boiss amp Huet Fl Orient 1 161 1867Ind loc lsquoBoiss et A Huet in pl Arm Exs 1853rsquoLectotype (designated here) Armenia in umbrosissupra Keuprubachu inter Trapezuntem et Baibout5500ndash6000 psm Maio 1853 A Huet du PavillonPlantae Orientales exsiccatae (G)

= C pectinata var adzharica Albov Prodr Fl Colchic19 1895Ind loc lsquoAdzharia Yaiumlla Medzybna prope pagumAkria 1500ndash2000 m (N A 1893 n 80)rsquoLectotype (designated here) Spuskrsquo srsquo yaily Medzybnykrsquo sel Akria 1500ndash2000 m 10 juin 1893 N AlboffPlantes drsquoAdjarie no 80 (G)

= C parviflora f latifolia O E Schulz Bot JahrbSyst 32 484 (1903)Ind loc lsquoad mare Caspium pr Lenkoran leg C AMeyer 1829 (H Boiss HPAc)rsquoLectotype (designated here) ad m Caspium 1829C A Meyer Enum Cauc no 1594 (G)

Description Biennial or rarely annual herb Height ofstem from base to the lowest peduncle of flower or fruit(6ndash) 7ndash27 (minus35) cm Stem erect flexuous oftenbranched from the base or simple glabrous Rosetteleaves shrivelled by anthesis pinnate sparsely hairywith two to four pairs of petiolulate circular-ovatelobate or sometimes pinnatipartite leaflets Caulineleaves (3ndash) 4ndash10 (minus11) auriculate pinnate or some-times pinnatisect above with 5ndash9 (minus11) denticulate tolobed or pinnatipartite leaflets Terminal leaflets(07ndash) 09ndash37 (minus48) times (02ndash) 03ndash16 (minus18) cm longerthan lateral leaflets petiolulate or sessile lanceolateLateral leaflets (05ndash) 06ndash24 (minus28) times 02ndash13(minus15) cm petiolulate or sessile lanceolate to ovateInflorescence racemose with 0ndash8 (minus9) lateral inflores-cences Sepals oblong (16ndash) 18ndash28 (minus30) times (05ndash)06ndash10 (minus11) mm margin membranous petals (31ndash)34ndash58 (minus64) times (10ndash) 11ndash22 (minus24) mm white oblan-ceolate apex rounded Stamens 6 anthers greenndashyellow filaments (20ndash) 23ndash36 (minus41) mm long stigmanot conspicuous Pedicels erect-patent and siliquaeexpanded and often secund


Ecology Moist sites in shady forests mixed or conif-erous forests from 50 to 3000 m (Iran)

Distribution area Mountain ranges in the BalkanPeninsula (Bulgaria Greece) northern Turkey the

Caucasus region and northern Iran A single recordfrom the Hatay province in southern Turkey

ACKNOWLEDGEMENTS

We would like to thank our colleagues from BulgariaM Anchev S Stoyanov Turkey S Hayirlioglu-AyazM Keles Russia D Geltman V G Onipchenko andSlovakia M Kolniacutek for their help and accompany-ing during the fieldworks curators of herbaria givenin the Material and Methods section for allowing usto study herbarium specimens and finally K Trem-etsberger Vienna Austria for the help in molecularlaboratory This study was suported by the ScientificGrant Agency of the Ministry of Education of the Slo-vak Republic and the Slovak Academy of SciencesVEGA Bratislava (grant no 3042) by the Researchand Development Support Agency (grant no APVT-51-026404) and by the Ministry of Education Youthand Sports of the Czech Republic (grant no0021620828)

REFERENCES

Adams JM 1997 Global land environments since the lastinterglacialURL httpwwwesdornlgovernqennerchtml[accessed on 30 September 2005) Oak Ridge TennesseeOak Ridge National Laboratory

Albach D Martiacutenez-Ortega MM Chase MW 2004 Veron-ica parallel morphological evolution and phylogeography inthe Mediterranean Plant Systematics and Evolution 246177ndash194

Al-Shehbaz IA 1988 The genera of Arabideae (CruciferaeBrassicaceae) in the southeastern United States Journal ofthe Arnold Arboretum 69 85ndash166

Anchev ME Goranova V 1999 Reports (1031ndash1049) InKamari G Felber F Garbari F eds Mediterranean chro-mosome number reports 9 Flora Mediterranea 9 331ndash339

Applied Biosystems 1996 AFLPtrade plant mapping protocolFoster City CA PE Applied Biosystems A Division ofPerkin-Elmer

Assenov I 1970 Cardamine L In Jordanov D ed FloraRepublicae Popularis Bulgarica Vol 4 Sofia In AedibusAcademiae Scientarum Bulgaricae 418ndash446

Candolle AP 1821 Regni vegetabilis systema naturale Vol 2Paris Treutell et Wuumlrz

Cullen J 1965 63 Cardamine In Davis PH ed Flora of Tur-key and the East Aegean Islands Vol 1 Edinburgh Univer-sity Press 438ndash444

Denk Th Frotzler N Davitashvilli N 2001 Vegetationalpatterns and distribution of relict taxa in humid temperateforests and wetlands of Georgia (Transcaucasia) BiologicalJournal of the Linnean Society 72 287ndash332

Doyle JJ Doyle JL 1987 A rapid DNA isolation procedurefor small quantities of fresh leaf tissue Phytochemical Bul-letin of the Botanical Society of America 19 11ndash15

188 J KUBERA ET AL


Dreyer LL Jordaan M 2000 Brassicaceae (Cruciferae) InLeistner OA ed Seed plants of southern Africa families andgenera Strelitzia 10 184ndash191

Everitt B 1986 Cluster analysis 2nd edn (Reprint 2) NewYork Gower Halsted Press

Fineschi S Taurchini D Grossoni P Petit RJ Ven-dramin GG 2002 Chloroplast DNA variation of whiteoaks in Italy Forest Ecology and Management 156 103ndash114

Franzke A Hurka H 2000 Molecular systematics andbiogeography of the Cardamine pratensis complex (Bras-sicaceae) Plant Systematics and Evolution 224 213ndash234

Gilibert JE 1782 Flora Lithuanica inchoata Vol 1 GrodnoTypis SRM

Gohil RN Ashruf M Raina R 1981 Cytotaxonomicalconspectus of the flora of Kashmir II Chromosome num-bers of 51 dicotyledonous species Herba Hungarica 20343ndash49

Grossgeim AA 1950 Rod 444 Cardamine L In GrossgeimAA ed Flora Kavkaza Vol 4 2nd edn Moscow LeningradIzdatelrsquostvo Akademii Nauk SSSR 181ndash183

Hampe A Arroyo J Jordano P Petit RJ 2003 Rangewidephylogeography of a bird-dispersed Eurasian shrub con-trasting Mediterranean and temperate glacial refugiaMolecular Ecology 12 3415ndash3426

Hedge J 1968 Cardamine In Rechinger KH ed Flora Iran-ica Flora des iranischen hochlandes und der umrahmendengebirge 57 (Cruciferae) Graz Akademische Druck-u Verlag-sanstalt 193ndash197

Heuertz M Fineschi S Anzidei M Pastorelli R SalviniD Paule L Frascaria-Lacoste N Hardy OJ Veke-mans X Vendramin GG 2004 Chloroplast DNA varia-tion and postglacial recolonization of common ash(Fraxinus excelsior L) in Europe Molecular Ecology 133437ndash3452

Hewitt GM 2002 The genetic legacy of the Quaternary iceages Nature 405 907ndash913

Hewitt GM 2004 Genetic consequences of climatic oscilla-tions in the Quaternary Philosophical Transactions of theRoyal Society of London Series B 359 183ndash195

Holmgren PK Holmgren NH Barnett LC 1990 IndexHerbariorum Part I The Herbaria of the World 8th ednRegnum Vegetabile 120 1ndash693

Jafri SMH 1973 Brassicaceae In Nasir E Ali SI eds Floraof west Pakistan Vol 55 Karachi Herbarium Departmentof Botany University of Karachi

Jalas J Suominen J 1994 Atlas florae Europaeae Vol 10Helsinki The Committee for Mapping the Flora of Europeand Societas Biologica Fennica Vanamo

Jee V Dhar U Kachroo P 1983 Chromosome numbers ofsome alpine-subalpine taxa of Kashmir ndash Himalaya HerbaHungarica 222 23ndash36

Jones BMG Akeroyd JR 1993 40 Cardamine L In TutinTG Heywood VH Burges NA Valentine DH Walters SMWebb DA eds Flora Europaea Vol 1 2nd edn Psilotaceaeto Platanaceae Cambridge Cambridge University Press346ndash352

Ketskhoveli NN 1979 Flora Gruzii Vol 5 2nd edn TbilisiMetsniereva

Khatri KS 1988 Studies in the Caucasian species of Car-damine L Phyton 28 55ndash80

Kimata M 1983 Comparative studies on the reproductivesystems of Cardamine flexuosa C impatiens C scutata andC lyrata Cruciferae Botanical Magazine Tokyo 96 299ndash312

King RA Ferris C 1998 Chloroplast DNA phylogeography ofAlnus glutinosa (L) Gaertn Molecular Ecology 7 1151ndash1161

Kit Tan 2002 16 Cardamine L In Strid A Kit Tan edsFlora Hellenica Vol 2 Ruggel ARG Gantner Verlag KG178ndash184

Klecka WR 1980 Discriminant analysis (Sage UniversityPapers Series Quantitative applications in the social sci-ences no 19) Beverly Hills CA Sage

Krebs P Conedera M Pradella M Torriani D Felber MTinner W 2004 Quaternary refugia of sweet chestnut(Castanea sativa Mill) an extended palynological approachVegetation History and Archaeobotany 13 145ndash160

Krzanowski WJ 1990 Principles of multivariate analysisOxford Clarendon Press

Kuddddera J Valko I Marhold K 2005 On-line database of thechromosome numbers of the genus Cardamine (Brassi-caceae) Biologia (Bratislava) 60 473ndash476

Lakutttticccc D Novddddicccc R Kuddddera J Marhold K 2006 Car-damine pancicii (Brassicaceae) a neglected species of theBalkan Peninsula morphological and molecular evidenceWilldenowia 36(Special issue) 177ndash191

Lanjouw J Stafleu FA 1957 Index Herbariorum Part II (2)Regnum Vegetabile 9 1ndash295

Legendre P Legendre L 1998 Numerical ecology Englishedition 2nd edn Amsterdam Elsevier

Lihovaacute J Marhold K 2006 Phylogenetic and diversity pat-terns in Cardamine (Brassicaceae) ndash a genus with conspicu-ous polyploid and reticulate evolution In Sharma AKSharma A eds Plant genome biodiversity and evolutionVol 1C Phanerogams (angiosperm ndash dicotyledons) EnfieldNH Science Publishers Inc 149ndash186

Lihovaacute J Marhold K Neuffer B 2000 Taxonomy of Car-damine amara (Brassicaceae) in the Iberian PeninsulaTaxon 49 747ndash763

Lihovaacute J Marhold K Tribsch A 2003 The Cardaminepratensis (Brassicaceae) group in the Iberian peninsulataxonomy polyploidy and distribution Taxon 52 783ndash802

Lihovaacute J Marhold K Tribsch A Stuessy TF 2004a Mor-phometric and AFLP reevaluation of tetraploid Cardamineamara (Brassicaceae) in the Mediterranean SystematicBotany 29 134ndash146

Lihovaacute J Shimizu KK Marhold K 2006 Allopolyploid ori-gin of Cardamine asarifolia (Brassicaceae) incongruencebetween plastid and nuclear ribosomal DNA sequencessolved by a single-copy nuclear gene Molecular Phylogenet-ics and Evolution 39 759ndash786

Lihovaacute J Tribsch A Stuessy TF 2004b Cardamine apen-nina a new endemic diploid species of the C pratensis group



(Brassicaceae) from Italy Plant Systematics and Evolution245 69ndash92

Loumlvkvist B 1956 The Cardamine pratensis complex Outlineof its cytogenetics and taxonomy Symbolae Botanicae Upsa-lienses 142 1ndash131

Loumlvkvist B 1957 Experimental studies in Cardamine amaraBotaniska Notiser 110 423ndash441

Maacutejovskyyyy J et al 1974 Index of chromosome numbers ofSlovakian flora Part 4 Acta Facultatia Rerum NaturaliumUniversitatia Comenianae Botanica 23 1ndash23

Marcussen T 2003 Evolution phylogeography and taxon-omy within the Viola alba complex (Violaceae) Plant Sys-tematics and Evolution 237 51ndash74

Marhold K 1992 Multivariate morphometric study of theCardamine amara group (Cruciferae) in the Carpathiansand Sudeten mountains Botanical Journal of the LinneanSociety 110 121ndash135

Marhold K 1994a Taxonomy of the genus Cardamine L(Brassicaceae) in the Carpathians and Pannonia I Car-damine pratensis group Folia Geobotanica Phytotaxonomica29 335ndash374

Marhold K 1994b Chromosome numbers of the genus Car-damine L (Cruciferae) in the Carpathians and in PannoniaPhyton (Horn Austria) 34 19ndash34

Marhold K 1995 Taxonomy of the genus Cardamine L(Brassicaceae) in the Carpathians and Pannonia II Car-damine amara L Folia Geobotanica Phytotaxonomica 3063ndash88

Marhold K Huthmann M Hurka H 2002 Evolutionaryhistory of the polyploid complex of Cardamine amara (Bras-sicaceae) isozyme evidence Plant Systematics and Evolu-tion 233 15ndash28

Moench C 1794 Methodus plantas horti botanici et agrimarburgensis a staminum situ describendi MarburgiCattorum In officina nova libraria academiae

Muumlller OF 1777 Flora danica Vol 513 KoslashbenhavnTrykt hos Hans Kogl Majest Hof-Bogtrykker NicolausMuller

Muriacuten A 1960 Substitution of cellophane for glass covers tofacilitate preparation of permanent squashes and smearsStain Technology 35 351ndash353

Nei M Li WH 1979 Mathematical model for studying geneticvariation in terms of restriction endonucleases Proceedingsof the National Academy of Sciences of the USA 76 5269ndash5273

Pernyyyy M Tribsch A Anchev ME 2005b Infraspecific dif-ferentiation in diploid Cardamine acris (Brassicaceae) fromBalkan Peninsula morphological and molecular evidenceFolia Geobotanica 39 405ndash429

Pernyyyy M Tribsch A Stuessy TF Marhold K 2005aTaxonomy and cytogeography of Cardamine raphanifoliaand C gallaecica (Brassicaceae) in the Iberian PeninsulaPlant Systematics and Evolution 254 69ndash91

Petit RJ Aguinagalde I de Beaulieu J-L Bittkau ChBrewer S Cheddadi R Ennos R Fineschi S GrivetD Lascoux M Mohanty A Muumlller-Starck GDemesure-Musch B Palmegrave A Martiacuten JP Rendell SVendramin GG 2003 Glacial refugia hotspots but not

melting pots of genetic diversity Science 300 1563ndash1565

Podani J 2001 SYN-TAX 2000 Computer programs for dataanalysis in ecology and systematics Userrsquos manual Budap-est Scientia Publishing

Rollins RC 1993 The Cruciferae of continental North Amer-ica Stanford CA Stanford University Press

SAS Institute 2000 SAS OnlineDocreg Version 8 (availableonline) Cary North Carolina SAS Institute

Schneider S Roessli D Excoffier L 2000 Arlequin vers 2000 a software for population genetics data analysisGeneva Genetics and Biometry Laboratory University ofGeneva

Schols P Dessein S Drsquohondt C Huysmans S Smets E2002 Carnoy a new digital measurements tool for palynol-ogy Grana 41 124ndash126

Schoumlnswetter P Tribsch A Barfuss M Niklfeld H 2002Several Pleistocene refugia detected in the high alpine plantPhyteuma globulariifolium Sternb amp Hoppe (Campanu-laceae) in the European Alps Molecular Ecology 11 2637ndash2647

Schoumlnswetter P Tribsch A Stehlik I Niklfeld H 2004Glacial history of high alpine Ranunculus glacialis (Ranun-culaceae) in the European Alps in a comparative phylogeo-graphical context Biological Journal of the Linnean Society81 183ndash195

Seymour FC 1969 The flora of New England RuthlandVermont The Charles E Tuttle Co

Sneath PH Sokal RR 1973 Numerical taxonomy Principlesand practice of numerical classification San Francisco CAWH Freeman

Stoyanov N Stefanov B 1948 Flora na Bulgariya 3rd ednSofia Universitetska Pechatnica

Swofford DL 2003 PAUP Phylogenetic analysis using par-simony (and other methods) Version 4 Sunderland MASinauer Associates

Trautvetter ER 1876 Plantarum messes anno 1874 inArmenia a Dre G Radde et in Daghestania ab A Becker fac-tas commentatus est E R a Trautvetter Trudy Imperator-skago S-Petersburskago Botanicheskago Sada 4 (1) 97ndash192

Urbanska KM Hurka H Landolt E Neuffer B Mummen-hoff K 1997 Hybridization and evolution in Cardamine(Brassicaceae) at Urnerboden Central Switzerland biosys-tematics and molecular evidence Plant Systematics andEvolution 204 233ndash256

Vos P Hogers R Bleeker M Reijans M van de Lee THornes M Frijters A Pot J Poleman J Kuiper MZabeau M 1995 AFLP a new technique for DNA finger-printing Nucleic Acids Research 23 4407ndash4414

Willis KJ 1994 The vegetational history of the BalkansQuaternary Science Reviews 13 769ndash788

Yeh FC Yang R-C Boyle TJ Ye Z-H Mao JX 1997 POP-GENE the user-friendly shareware for population geneticanalysis Edmonton AB Molecular Biology and Biotechnol-ogy Centre University of Alberta

Zohary M Heyn CC Heller D 1980 Conspectus floraeorientalis Fasc 1 Jerusalem The Israel Academy of Sci-ences and Humanities

190 J KUBERA ET AL


APPENDIX

Survey of herbarium specimens of C impatiens(from the area of sympatry with C pectinata) andC pectinata (reflecting the whole distribution area ofthis species) from herbaria B BM G K KW LE LEUP PR PRC SAV SO SOM W WU

CARDAMINE IMPATIENS LBULGARIA STARA PLANINA in silvaticis m Ruj nonprocul pagum Odorovci 16vi1903 Iv K Urumov(SOM 32080) Odorovci Iv K Urumov (PRC) propeTeteveni 6vi1910 Iv K Urumov (SOM 32048) in sil-vaticis ad pagum Glo|ene distr Pleven 14vi1910 IvK Urumov (SOM 32049) bei Trojan 1900 Iv K Uru-mov (PRC) Trojan Balkan 1899 Iv K Urumov (PRC)Trojan Balkan 22vii1898 Iv K Urumov (WU)Gabrovo dolina r Panicharka u Khadisi Tsoneva mostasredi skal 29v1930 A Yurkovskii (SOM 32036)Gabrovo po kamenisti mesta v dolinata na r Pani-charka krai Khadzhi Tsoneviya most 29v1930 AYurkovskii (SO 83221) Petkovo Niva Gabrovo25iv1897 Iv Neuchev (SOM 32073) in saxosis etpratis mt Balkan central ad riv Star ad v Gabrovo450 m 3v1907 B Davidov (SOM 32062) in saxosis etpratis mt Balkan central ad riv Star ad v Gabrovo400 m 2vi1907 B Davidov (SOM 32067) Kotkov pgTumla 19v1891 K Polaacutek (PRC) Chernomorskokraibrezhie reservat lsquoKamchiyarsquo do Lesinski azmak9v1996 D Stoyanov amp D Ivanov (SO 98398) VITO-

SHA Ad Bojana prope Sofia 17v1910 Iv K Urumov(SOM 32055 32052 SO 83220) Vitosha po ptya Kopi-toto-Molina skala 25v1964 B Kitanov (SO 28210) inMt Vitota 1889 Tkorpil (PRC) LOZENSKA PLANINALozenska planina in declivibus umbrosis infer fruticessupra pagum Dolni Losen distr Sofia 7vi1978 NVihodcevski (SOM 139181) PLANA In saxosis humidismt Plana ad faucem rivi Iskar 750 m 3v1910 BDavidov (SOM 32066) OSOGOVSKA PLANINA In uligi-nosis umbrosis ad p Skakavica distr Kjustendil1902 Iv K Urumov (SOM 32057 32054) BELASITSA

PLANINA Belasitsa planina po ptya za vrkh Tumba(portala s Skrt) 24v1998 S Kostaninova (SO 99754)Belasitsa iz vlazhnite dolove nad s Kolarovo19v1999 P Zhelev amp G Gogushev (SOM 154937)RILA in silvaticis humidis mt Manastirska Rila adcoenobium Rila 1200 m 21v1912 B Davidov (SOM32065) Rila pl pod Musala do r Bistritsa viii1897 VStribrni (SO 28202) Rila planina in silvis Piceae excel-sae umbrosis inter Saparevo et Skakavica 1120 m31vii1923 A Novaacutek 225 (PRC) Skakavica planina10vii1902 Iv K Urumov (PR) in silvaticis saxosismt Malka Rila Titman vrch versus rivi Iskar 1150 m27v1911 B Davidov (SOM 32070) RODOPI Ad Jun-dola m Rhodope 1912 Iv K Urumov (SOM 32046)

Sredni Rodopi Trigradsko zhdrelo pokrai ptya podpeschlivi kamenisti mesta 15vi1986 M Anchev(SOM 145068) M Rhodope in valle fl Berna supraRajkovo 11viii1961 G Zochev (SO 28209)GREECE NORTH-EAST Nom Pellis Ep AlmopiasMt Tzena N of the village of Notia by a forest road1600ndash1750 m 41deg09primeN 22deg11primeE 30v1999 Strid et al49012 (G) NORTH CENTRAL NW Macedonia Pisode-rion damp shady place in ravine near the Hagia TriasMonastery 4000 ft 28vi1932 A H G Alston amp N YSandwith 1037 (K) Flora of Thrace Xanthia-ShaninRoad 600 ft 23v1933 H G Tedd (K) EAST CEN-

TRAL Olymp Eichenwaumlldern im Enipcostal 1400ndash1600 m vii1938 E Behr (W 21519)TURKEY A5 KASTAMONU Prov Kastamonu IlgazDa 5000 ft 9vi1954 Cullen 1963 (K) A7 GIRESUNProv Giresun Tamdere Yavuzkemal near Karincaby roadside 1500 m 13viii1952 Davis et al 20710(BM W 8710 K) A7 TRABZON Maccedilka HamsikoumlyZigana da Bachufer 1400 m 18vii1958 A Huber-Morath 14788 (G) A7 GUumlMUumlSHANE Armenia TurcicaSzanschak Guumlmuumlschkhane Kurzul im dumetis6vi1894 C Haussknecht 5754 Iter Orientale 1894(PRC PR G) A8 RHIZE Prov Rize Ikizdere-Carankaya yayla 1000 m 25viii1952 Davis 20886 ampDodds (BM) Bords des torrents de la valleacutee sous-alpine de Djimil (Lazistan) vers 2000 meters drsquoalti-tude 1866 B Balansa (PRC K G) A8 ARTVIN ProvCoruh (Artvin) Kardevan dag (Yalnizccedilan-Da lari)near Kuumltuumll yayla 2200 m 28vi1957 Davis 30240 ampHedge (K G) idem 2100 m 18vi1957 Davis 30203amp Hedge (K) B9 SATAK Prov Van distr SatakKavuccedilsahap Dag 6500 ft 22vii1954 Davis 23011 ampO Polunin (BM K)RUSSIA KRASNODARSKII KRAI Caucasus occidentalisdistr Lazarevskoe in valle fluminis Shakhe infravicum Solokhaul 300 m 26vi1979 V Vataacutek (W05699) Chernomor okr Sochi 28v1895 W Lipsky(LE) Caucasus occidentalis distr Sochi in valleaugusta rivi Agva ad cataractus lsquoOrekhovye vodopadyrsquodicta c 300 m 11vi1978 V Vataacutek amp A V zda (W05697) Caucasus occidentalis distr Khosta reserva-tum naturale lsquoSanshitovaya roscarsquo 100ndash130 m16vi1978 V Vataacutek amp A V zda (W 05698) ZapadniiKavkaz Krasnaya polyana g Achishkho bukovyi les5vii1988 [sine coll] (LE) Zapadne Zakavkazrsquoe gAchishkho JZ sklon 1850 m bukovyi les 3vii1979I A Sakhalova amp A A Pautov (LEU) Chernomor-skaya g Krasnaya Polyana Yu sklon Achishkho30vi1915 A Abletsov (LE) Krasnodarskii krai Lab-inskii r-n okr st Khodzrsquo 26v1989 Dolmatova et al(LE) Severnii Kavkaz Krasnodarskii krai Mos-tovskoi r-n pos Psebai okolo sel Nikitovka pravyibereg r Malaya Laba 27vii1990 Yu Menitskii et al(LE) Severo-Zapadnii Kavkaz vodorazdel r BolrsquoshayaLaba 23vii1930 A I Leskov (LE) ADYGEA Severnii

g

g

g

e

e



Kavkaz okr Maikopa 10vi1923 N L Pastuchov(LE) KARACHAY-CHERKESSIA Severnii KavkazStavropolrsquoskii krai okr sel Leso-Kefarrsquo 16 km v verkhpo tech r Kefarrsquo pravyi bereg 10viii1989 YuMenitskii et al (LE) Teberdinskii zapovednik posDombai les 15ix1977 A O Malunaev 77-931 (LE)Teberda 4300 ft 13vi1904 I Ya Akinfieva (LE) rTeberda 22vii1939 V L Komarov amp I V Komarova(LE) Teberda 1939 V L Komarov amp I V Komarova(LE) Kuban prov levyi bereg reky Teberdy 4200 ft13ndash16vi1908 N Busch (LE) Kuban prov Teberda4200 ft 14vi1907 E A Endaurova (LE) KarachayKhumarinskii okrug lugovina po r Teberde mezhduMikoyan-Shakharom i aulom Senty 13vii1928 K IMarusyak (LE) Karachevo-Cherkesskaya avtonom-naya oblastrsquo Stavropolskii krai Dombai oddelenieTeberdinskogo zapovednika 7ix1974 M Kotov(KW) Stavropolrsquoskii krai Karachaevo-CherkesskayaAO 6ndash10 km k vostoku ot Ust-Dzheguty dolinnyi lespo r Dzhegonas 18vi1988 Geltman et al (LE)Kuban obl Batalyatinsk 2vi1892 W Lipsky (LE)Karachaevo-Cherkesskaya AO 7 km k vostoku ot sCherkeska mezhdu s Markoli i Nizhnaya Mara9vii1989 Geltman et al (LE) KABARDINO-BALKARIYA Kavkaz Kabardino-Balkarskii zapoved-nik Tserek Balkarskii verkhoviya r Kara-su10vii1963 Tikhonova (LE) Terskaya obl selNalrsquochik 2v1970 V Palrsquotseva (LE) Kab-Balk ASSRGolubye ozera lug na pravom beregu r Cherek u 1-ogo ozera 9vi1964 E Dobynina (LEU) NORTH

OSSETIYA Severnaya Ossetiya teritoriya Severo-Osetinskogo zapovednika verkhovrsquoya r Mamison-donushchelrsquoe Zemagon 29viii1989 Yu Menitskii et al(LE) Caucasus Ossetia districtus Alagir 320ndash330 m20v1861 Ruprecht (LE) Caucasus Ossetia pr Ala-gir 14iv1899 Marcowitsch (LE) Severo-Osetinskiigos zapovednik i ego okhrannaya zona Tseiskoelesnichestvo levyi bereg reky Tseidon nizhe turbazyTsei 8vii1987 B P Serov amp I V Sokolova (LE)Kavkaz Ossetia Vladivkavkaz near Lisaya mt14vi1900 G A Levitskii (KW) STAVROPOLrsquoSKII KRAIFlora Stauropolensis 21vi1881 A Normann (LE)Stavropol 1889 A Normann (LE) Stavropolrsquoskayavozvyshennostrsquo shirokolistvennye lesa po vossklonam g Strizhament k severu od st Novo-Ekat-erinovskaya 24vi1988 Geltman et al (LE) KavkazZheleznovodsk v1887 I Akinfiev (LE) Terskayaoblastrsquo Zheleznovodsk (u vershiny g Zheleznaya) vlesu u dorogy ne chasto 24vi1906 P Tischevsky(KW) Prov Terek distr Pyatigorsk gora Beshtau17vii1913 A Gordiagin (LE) CHECHENIYA Chech-eno-Ingushetiya SSR Sovetskii raion dolina rekyChanty-Argun okr sel Itum-Kale na beregu ruchrsquoya26vi1967 V M Prima (LE) Checheno-IngushetiyaSSR Groznenskii raion okr stanitsy Petropav-lovskaya opushka lesa 14vi1967 V M Prima (LE)

DAGESTAN Dagestan subalpinskii poyasrsquo mezhduKodorskim perevalom i s Khupro 17vii1904 NBusch (LE) Daghestan subalpiiskii poyasrsquo mezhdu sBezhita i perevalom Michishlrsquo 12vii1904 N Busch(LE) Tsyarashinskii rn dolina Avarskno Keieumezhdu s Sovetskoe i Tsyarasha v 7 km yuzhneKosob-orta (1400ndash1500 m) vlazhnoi galernik19vii1961 N N Tsvelev et al (LE) DagestanskayaASSR raion Gumbetovskii mezhdu sel Burtunai iperevalom k sel Danukh bukovyi les po sklonu krechke 1761961 N N Tsvelev et al (LE) Dagestan-skaya ASSR raion Gumbetovskii mezhdu sel Bur-tunai i perevalom k sel Danukh zasurennii lug17vi1961 N N Tsvelev et al (LE) Prov Dagestandistr Dargi pr Murguk 2700 ft in humidis ad rivu-lum 7vii1898 Th Alexeenko (LE) Prov Dagestandistr Temir-Chan-Schura [Buinaksk] in declivitateboreali m Madygir-tau supra Erpeli ad rivulum4100 ft 29vi1898 Th Alexeenko (LE) Prov Dag-estan Temir-Chan-Schura [Buinaksk] T Ch SchuraErpeli 1700ndash1800 ft 19vii1897 Th Alexeenko (LE)Dagestan Gunibskii okrug verkhoviya Kara-Lagur2400 m 25viii1928 leg A Poretskii (LE) Dagestan-skaya ASSR Buinakskii raion vost sklon Gimrin-skogo khr sel Verkhni Kazanishi 7vi1961 N NTsvelev et al (LE) Prov Dagestan distr Samur locoZassyl-dere inter Lutszek et Kina in Pinetis 6000ndash6600 ft 9viii1898 Th Alexeenko (LE) Dagestan-skaya ASSR Akushinskii raion mezhdu sel Garshinai Akusha dolina r Akusha 9vii1961 N N Tsvelevet al (LE) Dagestanskaya ASSR Rutulrsquoskii r-nsel Khmot mezhdu s Akhty i s Rutul Ushcheloepravogo pritoka r Samur 27vi1961 N N Tsvelevet al (LE)GEORGIA ABKHAZIA Abkhazskaya ASSR Gogrskiiizvestnyakovyi massiv yuzhnii makrosklon khr Ber-chilrsquo bas r Sandripsh okr p Gyuzlya 12 km k SV OTg Gagra 27vi1989 Geltman et al (LE) Abkhaziyaverkhnaya chastrsquo levogo borta r Bzibrsquo gora Adats25viii1933 Yu R Roskov (LE) Abkhazskaya ASSRGruzinskaya SSR 25 km above Ritsa lake Ritsa onacid rock at lumber camp in mixed coniferousdecidu-ous forest Lashipse-Bzib valley 24vi1971 C RFrazer-Jenkins 2756 (BM) Caucasus occidentalisdistr Gudauta in vicinitate lacus Rica 1000ndash1300 m21vi1977 V Vataacutek (W) Caucasus occidentalis distrGudauta in vicinitate lacus Mzymt et angustiarumAcetuki 1700ndash2300 m 16vi1971 V Vataacutek Iter Cau-casicum 1971 (PR) Caucasus occidentalis Georgiadistr Sukhumi vicinitas Latior pagi Pskhu locoSanchava dicto secum rivum Akhei 700ndash800 m5viii1983 V Vataacutek (W) Caucasus occidentalis distrGulripsch in vicinate pagi Sakeni 1000ndash1100 m12vi1971 V Vataacutek Iter Caucasicum 1971 (PR)AJARIA Batumskaya obl Adzhariya bl st Nonusrsquo-ogly 1300 m 25vi1913 D Litvinovrsquo (LE) GURIA

192 J KUBERA ET AL


Caucasus occidentalis distr Chokhatauri vicinitaspagi Bakhmaro montes Meskhadski khrebet indeclivibus septentrionalibus montis Sakornia 2000ndash2400 m 20vii1980 V Vataacutek (W 05702) SAMTSKHEAdugenskii r-n okr Abastunani 9viii1977 YuMenitskii (LE) pr Borshom vi G Radde 134 (LE)Borshom vii1865 G Radde (LE) Borshom 1884 GRadde (LE) Bakuriani 20vi1913 I Novopokrovskii(LE) prope Bakuriani vi1881 A I Brotherus amp V FBrotherus (LE BM) Prov Kartli (Krsquoartli Kartli) Tri-aletisches Gebirge (Trsquorialetrsquois krsquoedi Trialetis kedi)Nordhang des Berges Kokhta gora (Kochta gora) c2 km ouml Bakuriani 1800ndash1900 m 43deg32primeE 41deg45primeN15vii1997 M Staudinger (W 01005) Gruz SSRBorzhomskii r-n okr sel Tsikhirzhvari 1800 mbukovyi les 27vi1980 T Popova et al (LE) propeurb Tambow in silva umbrosa subhumida 5vi1902fl 22vi1902 fr I Schirajevski (LEU KW) KARTLIKavkaz Tiflis 27v1865 E Overin (LE) Caucasuscentralis ad fl Aragwa Lagowski (LE) Kudarskiiraion alrsquopiiskii lug na perevale Dzedo 1950 m8viii1928 E Busch amp N Busch (LE) pod perevalomVanuri-ztseg pastbishche Erdusta 2200 m8vii1930 E amp N Busch (LE) Bass Maloi LiakhvyShuatskhuri g Nekh 18vii1933 E Busch amp NBusch (LE) Shuatskhuri senokosnii subalrsquopiiskii lugna g Nekh 2200 m 9vii1930 E Busch amp N Busch(LE) Shuatskhuri bukovyi les na urochishche Galer-gevden na levom borte ushchelrsquoya r Bestauty-don2000 m 7vii1930 E Busch amp N Busch (LE)Verkhne Kabustinskoe ushchelrsquoe 7viii1929 E Buschamp N Busch (LE) Ermani Zaleznik r VerkhniiErmani 21vii1937 I I Abramov (W 3341) Yugo-Osetinskii gorno-lugovoi statsionar mestnostrsquoErmani 24vii1938 B A Shtakelrsquoberg (LEU)Yugo-Osetinskii gorno-lugovoi statsionar mestnostrsquoErmani 15viii1938 G P Kvaratskheliya (LEU)Yugo-Osetinskii gorno-lugovoi statsionar mestnostrsquoErmani levyi bort Sredne Ermanskogo ushchelrsquoyav bereznyake 10vii1938 I I Abramov (LE)subalrsquopiiskii senokosnii lug levogo borta VerkhnegoErmanskogo ushchelrsquoya 2550 m 18viii1929 EBusch amp N Busch (LE) Chaparukhskoe ushchelrsquoe lesna urochishche Zandzhiket bliz Lrsquobira 1900 m10viii1930 E Busch amp N Busch (LE) Kudarskiiraion gora Nasty-tsup (Mikhal-Gabyr) les 2000 mviii1928 E Busch amp N Busch (LE) MTIULETI InPasanaur 550 m 26v1861 E Overin (LE) U perev-ala Gudaurrsquo 17vii1893 W Lipsky (LE) KHEVSURETIKhevsuria subalp lugy mezhdu uroch Mirgva guba is Batsrsquoaligo 6vii1903 N Busch (LE) KAKHETISabuck 23v1820 [sine coll] (LE) Kakheti GrosserKaukazus (Kavkasionis kedi) naturreservat Lagode-khi (Lagodekhis nakrzsali) schluchtartiger Abschnittbis zum Wasserfall 550ndash900 m 46deg18prime12PrimeE 41deg52prime33PrimeN19v2001 G M Schneeweis amp A Tribsch (W 10200)

ARMENIA TAVUSH Idzsevanskii r-n Azartsii i sTekhut 1300 m 7vi1973 [sine coll] (BM) Tavushprovince Ijevan distr c 6 km N of Dilijan E ofAghartsin monastery 1460 m 44deg50primeE 40deg46primeNFagus orientalis and Quercus macranthera forest withsmall meadows and rocks in between 13vi2002 Fay-vush et al Optima Iter XI0371 (SAV) TranscaucasiaArmenia borealis distr Idzhevan prope p Idzhevanin declivibus monti Gatsha 10vi1937 C Gurvitsch(LE) GEGHARKUNIK Kotayk province Hrazdan distrvalley of river Marmarik c 18 km NW of Hrazdanc 1 km NW of village Artavaz 1940 m 44deg33primeE40deg38primeN dry to semihumid meadows tragacanth com-munity with Astragalus microcephalus 17vi2002Fayvush et al Optima Iter XI0839 (SAV) VAYOTS

DZOR Vayotsdzor province Vajk distr c 2 km NE ofJermuk 2115 m 45deg42primeE 39deg51primeN forest with Quer-cus macranthera and meadows 25vi2002 Fayvushet al Optima Iter XI2023 (SAV) SYUNIK Megrinskiir-n gradsko-dubskoi les vyshe pos Vardanidzor6vii1956 T V Egorova et al (LE)AZERBAIDZHAN XANLAR Khanlarskii r-nOKREST s Zurnabad 17vi1988 G Z Davirsquoyan(LE) Taluumlsch 1836 D Kavelin (LE) ZAQATALA distrZakataly prope pag Dzhory 25v1936 T Heidemanamp H Kadinov (BM) Flora Caucasi Gub Elisabeth-pol Dis Nucha ad fl Jatuch-czai (Bum-czai) adripam 6900 ft 23vii1899 Th Alexeenko (LE) QUBAFlora Caucasi Gub Baku Dis Kuba [Quba] ad pagChinalug [Xinaliq] (Ketisch Lechin) SW versusMadzigerdrich in angustiis 6900 ft 22viii1900Th Alexeenko (LE) Transcaucasia AserbaidzhanKubatenskii r-n selo Laza les 18vii1973 Gatinov(LE) LENKORAN Lenkoranskaya expeditsiyamezhdu s Lekin i s Ofanz 8vii1931 N Shiiinskii648 (LE)IRAN MAZANDARAN Mazanderan Sang-Deh 30 kmSE Poleacute-Seacutefid in Rocky mt 2500ndash3000 m 11vii1974Renz (W 6842) Mazanderan Nezva Kuh areabetween Orim and Taru in the forest 35deg59primeN53deg11primeE 2000 m 6vii1959 P Wendelbo (W 14903)

CARDAMINE PECTINATA PALL EX DC

BULGARIA STARA PLANINA prope Teteveni11v1910 Iv K Urumov [orig det as C graeca] (SOM32060) Etropole sd Iv K Urumov (PRC) KONIA-

VSKA PLANINA Koniavska planina po vlazhni mesta vdol severo ot Mrzen-Malinazha 21vii1957 D Jor-danov amp As Janev (SO 94144) RILA Rila pl iz elovitegori pokrai r Blagoevgradska Bistritsa 25vi1955 IvPenev (SO 28204) Zap Rila iz bukovite gory po sev-erniya sklon v mestnosta lsquoDobro polersquo 28vi1955 IPenev (SO 28206) Rila rezervat lsquoParangalitsarsquo vlazhnisenchesti mesta yugozapadnii sklon nad r Khaid-ushka 29vi1975 N Andreev (SOM 132165)



Zapadna Rila Partizanska polyana 21vi1957 IvBondev (SOM 109126) Rila pl nad monastira s dN Arnaudov (SO 28205) Rila iz gorata v raiona nalsquoPartizanska polyanarsquo iztochno ot Rilski monastir17vi1966 N Vihodcevsky (SO 28207) Wald zwischenBorowetz und Bistritza 13vi1964 W Hempel (B)Rila vlazhni mesta iz iglolistnata gora nad Borovets29vi1972 N Vihodcevsky (SO 30020) in silvisumbrosis m Rila 1909 Iv K Urumov (SOM 32079)in silvis umbrosis mt Tsarska Rila Sokolets 1300 malt 30v1910 B Davidov (SOM 32064) in silvaticishumidis mt Zarska Rila Zarska Bistriza 1350 m alt5vi1911 B Davidov (SOM 32043 SOM 32047 SO83223) in umbris silvarum M Rila viii1889 Velen-ovsky (PRC) in Mont Rila v1894 St iacutebrny (PRC)PIRIN Pirin rezervat lsquoBaiovi dupkirsquo vlazhni senchestimesta v sobchestva na biala mura 20vi1977 NAndreev (SOM 135641) Montes Pirin (centralis) adoppidum Bansko vallis Banderica c 1900 m solo cal-careo vii1936 I Klaacutettersky amp M Deyl (PR)GREECE SOUTHERN PINDOS Epirus prov Ioanninadistr Metsovo in latere boreali montis Kira ad austro-occidentem pagi Milea (secum viam ad Metsovoducentem) 1500 m 15viii1974 A Charpin et al (G106058) EAST CENTRAL Im m Olympo Thessaliaefagetis supra valle Stravits 1700ndash1900 m 23vii1937N Stoyanov amp D Jordanov (SOM 32037) in Pinetiumbrosis m Olympi Thesaliae 25v1851 de Hel-dreich (WU G) Thesalien Ossa-Gebirge SE Olymp1200 m 22deg42primeE 39deg50primeN 20v1989 H Malicky (W)in silvis fagetum supra Mon AG Dimitrios propepagum Csaigesi Ossa Mt (Thess) 26vii1936 OGrebenchikov (K)TURKEY A2 BURSA Bursa Mezit Deresi 23vi1976Y Akunza Queacutezel 6143 (G) Prov Kuumltahaya (Bursa)Tavsauli to Inegol 30 miles from Tavsauli 1200ndash1400 m 23vi1962 Dudley (W3496) A3-A4 BOLUBolu Ausar Ormani umgebung von Ausar Yaylasi ampSultankoumly Yaylasi c 1500 m 20v1970 I Bozakmanamp K Fitz 642 (W 13892) Woods above Abant Goumlluuml15vii1940 B V D Post (G) Seben dag bei BoluAbies bornmulleriana wald 12v1955 H Walter et al4556 (B) Prov Bolu Karadere to Yedigoumll bare banksin forest 650 m 18vii1962 Davis et al 37689 (K)Prov Bolu distr Ala Da 800 m 13viii1960 Khanet al 554 (K) A5 KASTAMONU Prov Kastamonu IlgazDag 5000 ft 9vi1954 Davis 21725 (BM) Prov Kas-tamonu distr Kastamonu Ilgaz-Da lari road Kasta-monu Ilgaz Abies forest 14viii1960 Khan et al 615(K) Kastamonu (Paphlagonien) IlgazndashKastamonuIlgaz Dag Abietum 2 km noumlrdlich der Passhoumlhe1700 m 21vi1955 A Huber-Morath 13762 (G) A5SINOP distr Ayancik Cangal Dag ab Ayancik 1100 m1vii1958 A Huber-Morath 14787 (G) A7 GIRESUN25 km S of Giresum 500 m 7vii1969 F Sorger (W8277) A7 TRABZON Trabzon 3 km S of Hamsikoumly

r

g

g

(c 45 km SSW of Trabzon) mixed forest of Picea andFagus with shrub layer of mainly Carpinus CorylusAlnus Sambucus Ribes and Rhododendron humidloamy soil c 1900 m vndashvi1959 Hennipman et al1867 (K B) Trabzon on slopes in woodland ascendingto Sumela Monastery c 53 km S of Trabzon 1180 m8v1988 C C Townsend 8883 (K) Prov TrabzonSoganli Pass growing with roots in running watervery bushy 7000 ft 17viii1962 P Furse 3985 (K)Prov Trabzon Soganli Pass 5000 ft 16viii1962 PFurse 3948 3966 (K) Trabzon Bayburt of 2 km Npass 2300 m 14vii1984 M Nydegger 19166 (G488601) in umbrosis supra Keuprubacku inter Tra-pezuntem [Trabzon] et Baibout [Baiburt] 5500ndash6000 m v1853 Huet du Pavillon Plantae Orientalesexsiccae (K) A8 RHIZE Prov Rize Ikizdere-Car-ankaya yayla 1000 m 25viii1952 Davis 20886 ampDodds (K) Regioacuten montagnese du Lazistan preacutes deDjimil 1800 m 19vii1866 B Balansa (P G) Evironde Rhizeacute bouddes chemins humides alt 400 m(Lazistan) vi1866 R du Parquet (BM) Environs deBehizeacute (Lazistan) sur les salus ombrageacutes des cheminscreux 27v1866 B Balansa (K PRC G) in alpibusLazistani A Eloy 113 (K) Rize 100 ft 22iv1959 KM Guichard TUR2459 (K) Senyuva Strasserand14iv1969 Steiner (W 04033) Rize SE oberhalb AyderIlica 16ndash18 km SE Ccedilamlihemsim 1400ndash1500 m8ix1980 F Sorger (W 8278) A8 ARTVIN Artvin byroadside flush 5 km S of side turning to DemirkentCcediloruh valley 50 km S of Artvin 10ix1970 C RFraser-Jenkins 2390 (BM) Prov Ccediloruh (Artvin)mountain above Artvin wet places in Picea forest bystream 1800 m 19vi1957 Davis 29775 amp Hedge (KW 8708) Prov Ccediloruh (Artvin) mountains aboveArtvin 1800 m 19vi1957 Davis 29775 amp Hedge(BM) Prov Artvin (A9) Yalniczam Gebirge bei Kar-akoumly oumlstlich von Savsat 1900 m im Felsgeroumlll einesBlockstromes in der oberen AbiesndashPicea-Nadel-waldstufe 5viii1981 Raus 4771 (B) 20 km E Savsatwald 2000 m 4viii1981 Raus (W8276) Artvin A9Yalniczam Silsilesi 8 km SE of Savsat c 1750 m10vii1967 Albury et al 3144 (K) Prov Ccediloruh(Artvin) Kordevan dag (YalnizccedilanndashDaglari) nearKuumltuumll yayla 2100 m Picea forest 18vi1957 Davis ampHedge 30203 (BM) A8 ERZURUM Erzeroum (Erzu-rum) M Zorab (Zohrab) [sd s coll] (K) B1 BALIKE-

SIR Anatolien Ccedilatal dag bei Balikesir 18vi1955 HWalter et E Walter 661 (B) C6 HATAY Plantae SyraeBorealis Montes Amanus valleacutes 800ndash3000 ft 1913M Haridjian 4670 (G)RUSSIA KRASNODARSKII KRAI Caucasus occidentalisdistr Adler (Sochi) in faucibus fluminis Psakho infravicum Kamenka 200ndash300 m 21vi1979 V Vataacutek (G2169346) Caucasus occidentalis Colchis distr Sochiin vicinitate pagi Izmailovka in valle fluminisMacesta 200ndash400 m 4vii1984 V Vataacutek (W) Cauca-

194 J KUBERA ET AL


sus occidentalis distr Sochi in vicinitate pagiMacesta ad ripam sinistram rivuli Macesta 50ndash200 m 7vi1978 V Vataacutek (G 269343) Flora Cauca-sica Zap Zakavkazrsquoe Khosta zapovednik 18vi1962N Spasskaya (LEU) L K Aleksandrova (LEU) ZapZakavkazrsquoe Krasnodarskii krai Khostinskii zapoved-nik syrie tenistye mesta 27vi1958 K K Smirnova(LEU) Caucasus occidentalis distr Sochi in faucibusrivi Agua (flumen Sochi 15 km ab ostio) 100ndash150 m17vi1979 V Vataacutek (W) Caucasus occidentalis distrSochi in valle augusta rivi Agva ad cataractuslsquoOrekhovye vodopadyrsquo dicta c 300 m 11vi1978 VVataacutek amp A V zda (G 269344) Yuzhnaya chast Cher-nomorskogo okruga mezhdu Adlerom i Tuanse25viii1929 S Ya Sokolov (LE) Krasnaya polyana pPudzik 6vii1936 N Busch (LEU) ZapadneZakavkazrsquoe gora Achishkho 1200 m bukovyi les3vii1979 T Onipko et al 238 (LEU) Chernomor-skaya Gub Achishkho 19vii1907 A N Kiritschenko(LE) Zapadnoe Zakavkazrsquoe Krasnaya polyana goraAchishkho sklon yuzhnoi ekspozitsii poyas bukovogolesa 4vii1985 I V Seferova (LE) ZapadneZakavkazrsquoe Yugo-zap sklon g Achishkho bukovyi les1800 m 3vii1979 L I Kurlakova (LE) Chernomor-skii okrug Kr Polyana yuzhni sklon g Achishkho1500 m 12vi1930 Yu P Byallovich (LE) Cherno-morg okr Krasnaya Polyana 3viii1895 (LE)Cherkesia Chernomorskii okr g Achishkho9vi1929 W Steup (LE) Chernomorskaya gub Ach-ishkho 19vii1907 A N Kiritschenko (LE) Caucasusoccidentalis distr Adler in vicinitate pagi KrasnayaPolyana vallis flumionis Mzymta 900ndash1000 m15vi1976 V Vataacutek (G 314654) ADYGEA ZapadniiKavkaz levyi bereg reky Belaya mezhdu selHamishki i Tuzerini Carpineto-Rhododendretum500 m 5vii1951 Ig V Vasiliev (LE) Kavkazskiizapovednik Maikopskii raion Tuzerini dolina rekyBelaya 3vii1935 Ig V Vasiliev (LE) ZapadniiKavkaz v 34 km ot Dakhovskoi (stanitsy) po levomuberegu reky Beloi suchoi kamenisty sklon s dubravoi500 m 5vii1951 Ig V Vasiliev (LE) Severo-ZapadniiKavkaz vodorazdel r Belaya Laba 16vii1930 A ILeskov (LE) KARACHAY-CHERKESSIA KavkazKubansk obl Teberda 4200 ft 1v1905 D Lumeinov(LE) Stavropolrsquoskii krai Pregradnenskii r-n pravyibort ushch r Bolrsquoshaya Laba v okr p Zagedan (40 kmk yu ot p Kurdzhinovo) 6vii1989 Dolmatova et al(LE)GEORGIA ABKHAZIA Abkhaziya okresnosti ozeraRitsa 22v1959 S Lipschitz (LE) Abkhazskaya ASRRitse-Abkharskii zapovednik u ozera Ritsa na bregu950 m 14v1955 M Kotov (KW) Zap ZakavkazrsquoeAbkhazskaya ASSR elovo-pichtovyi les SV sklon v10 km ot ozera Ritsa 1100 m 2vi1963 N G Avery-anova (LEU) Caucasus occidentalis distr Gagra apudlsquoGoluboe ozerorsquo in vicinitate rivi Bzyb 150 m

e

15vi1971 V Vataacutek Iter Caucasicum 1971 (PR)Zap Zakavkazrsquoe Avadkharskii zapovednik dorogana Avadkharu 12vi1957 A Bobrov (LEU) ZapZakavkazrsquoe Abkhazskaya ASSR r Avadkhara1900 m severnii sklon v tenistom i vlazhnom meste28vi1964 Belozub et al (LEU) Caucasus occidenta-lis distr Gudauta prope rivulum Avatchara 1700ndash2100 m 15vi1971 V Vataacutek Iter Caucasicum 1971(PR G) Caucasus occidentalis distr Gudauta in vic-inate lacus Mzymt et angustiorum Acetuki 1700ndash2300 m 16vi1971 V Vataacutek Iter Caucasicum 1971(PR) Abkhazskaya SSR Sukhumskii r-n ot Pskhu kAnchkho 12vii1980 T Popova et al (LE) Caucasusocidentalis distr Sukhumi in faucibus fluminisVostochnaya Gumista prope pagum Kaman 250 m23vi1977 V Vataacutek (G 289300) Caucasus occidenta-lis distr Sukhumi vicinitas pagi Akhalsheni in vallefluminis Zapadnaya Gumista 400ndash700 m 13vii1980V Vataacutek (W) Caucasus occidentalis distr Sukhumi indeclivibus apud rivum Gumista 10ndash150 m 13vi1971V Vataacutek Iter Caucasicum 1971 (PR) AbkhazskayaASSR Gulrsquoripshskii r-n okresnosti s Amtkel dolina rKholodnoi v 6 km k SSV ot Tsebelrsquody 25vi1989 ADolmatova et al (LE) Caucasus occidentalis distrGulripsh in vicinitate pagi Sakeni 1000ndash1100 m12vi1971 V Vataacutek Iter Caucasicum 1971 (PR) Cau-casus occidentalis distr Gulripsh in declivibus montisLachult in vicinitate pagi Sakeni 1300ndash2000 m10vi1971 V Vataacutek Iter Caucasicum 1971 (PR)AJARIA Prov Batum ad rupes humidas prope fortGoniiskii 1902 Alexeenko amp Woronow (LE) ProvBatum in silvis ad rupes littoralis prope fort Goniiskii100ndash300 ft 30v1902 Alexeenko amp Woronow (LE)Batumskii botanicheskii sad pri vkhode s Zel mysavmeste s Trachystemon i Petasites 28viii1987 ADmitrieva (LE) Batumskii botanicheskii sad2ii1962 A Dmitrieva (LE) idem 27v1962 A Dmi-trieva (LE) idem 16iv1973 A Dmitrieva (LE) idem19viii1980 A Dmitrieva (LE) idem 12ii1939 Popov(LE) Chernomorskaya Gub Sukhumskii okrugrsquoBatumskaya oblastrsquo vershina Aibgi 1911 I V Palibinamp N I Vorobev (LE) Kavkaz Batumsk obl i u okrBatuma na Zelenomrsquo mysu okrainy smeshan buko-vogo lesa 12vi1916 P N Krylov amp E I Shteinberg(LE) okr Batuma Zelenii mys listennyi les bliz botan-icheskogo sada 4vii1917 B Shishkin (LE) FloraCaucasica Chakva 3iv1896 B Levandowski (LE)Batumi po r Chakve na skalakh 21iv1950 M GPopov amp K D Stepanova (LE) Adzhariya okr selaKhulo 13ix1939 A Dmitrieva (LE) GURIAMakharadzevskii r-n doroga Chokhatauri-Kobuleti22iv1987 V Nikitin et al (LE) Zakavkazie GuriyaKutaisskaya gub Ozurgetskii u vindashvii1914 E IKikodse (LE) Caucasus occidentalis distr Chokha-tauri vicinitas pagi Bakhmaro montes Meskhadskikhrebet in declivibus septentrionalibus montis Sako-



rnia 2000ndash2400 m 20vii1980 V Vataacutek (B) Caucasusoccidentalis distr Chokhatauri ad orientem versuspagi Bakhmaro montes Meskhetski khrebet in decliv-ibus montis Mzis amosulis gori 2000ndash2300 m23vii1979 V Vataacutek amp G Esrandzhia Iter Caucasi-cum secundum 1979 (W) SAMTSKHE Transcaucasiamontes Ssuremenses praedium Rikoti [above Bor-jomi] 3900 ft 21vi1902 R Regel (LEU) Georgiainter Bakuriani et Tanbiszchuni vi1881 A I Broth-erus amp V F Brotherus (LE) Bakuriani Zagverivi1881 A I Brotherus amp V F Brotherus (BM) KARTLIles bliz Dzhalabet uroch Khusar 1300 m 27viii1928E Busch amp N Busch (LE) Shuatskhuri ushchelrsquoe rPotnisa bukovyi les 5vii1930 E Busch amp N Busch(LE) Bass Maloi Liakhvy Shuatskhuri Ikata bukovyiles 23vii1933 E amp N Bush (LE) verkhovrsquoya Kupt-inskogo ushchelrsquoya zarosl Rhododendron caucasicumna g Dzirisa 2100 m 8viii1930 E amp N Bush (LE)Chaparukhskoe ushchelrsquoe les Skatykom-kad 1900 m20vii1930 E Bush amp N Bush (LE) okr s Fatkudzhin1250 m 27viii1928 E Bush amp N Bush (K) KAKHETIKakheti Groser Kaukazus Natureserve LagodekhiSchluchtartiger Abschnitt bis zum Wasserfall 550ndash900 m 46deg18prime12PrimeE 41deg52prime33PrimeN 19v2001 G MSchneeweiss amp A Tribsch (WU)AZERBAIDZHAN QUBA Florae Trascaucasiae Gilavii1873 N Sredinsky (LE) LENKORAN Gub BakuTalysch in silvis umbrosis inter Sijon et Tazdy-Pok3300ndash4100 ft 3viii1897 Th Alexeenko (LE) Tran-scaucasia Azerbajdzhan prov Baku distr Lenkoraninter m Naduz-galasi et p Kagoj 14vii1929 L Pril-ipko (LE) ASTARA Azerb SSR Lenkoran okr Astara

r Ismi-su (Usmi-su) 20vii1975 Yu Menitskii ampT Popova (LE)IRAN GILAN Guilan vi1936 N Lindsay 1063 (BM)Qulan N Lindsay 870 (BM) Gilan m Talyschge-birge P Buhse (K) Prov Gilan in silvis supra Asa-lem (Navrud) ad viam versus Khalkhal (Herowabad)ducentem 200ndash800 m 15v1971 K H Rechinger (W11854 11855) MAZANDARAN Mazandaran in montissilvaticis 10 km merid Ab oppido Behsharh23vi1977 J Sojaacutek (SAV) Dimalu Mazenderan indense shade 23vii1940 W Koelz (W 6671) SarganBandar Gaz v1968 Sharif (W 6978) MazandaranKiasak Pete forest 1250ndash1600 m 10vii1974 Rense(W 6836) S of Chalus Caspian rain forest 500 ft27vi1962 P Furse 2885 (K) Elburz in foothills nearChalus 1500 ft 17iii1962 P Furse (K) Mazande-ran im Elbursgebirge bei Weyser oumlstlich des Chalus-Tales 1150 m 31viiminus6viii1968 E Steiner amp H MSteiner (W 2471) Mazanderan S of Amol c 60 m11vi1966 J C Archibald 2295 (K) Lajim forestManzanderan 3000 ft 20v1955 H F Mooney 6633(W 17400) Lajim forest Mazanderan under shadymoist calcareous clay bank in beech forest 3000 ft20v1955 H F Mooney (K) Ladjim Aghamashad(Fagetum) 800 m 30iv1965 Tregubov (W 1844)Ostan 2 entre Khozlok et Gurgan forecirct humide300ndash1600 m 6ndash7vi1956 F Schmidt 6022 (W24346G) Sargan Naharkhorran 21iv1974 Riedl ampErshad (W 21444) in montis Astrabadensis propeSiaret ivndashv1858 [without leg] (K G) Prov Gorgan(Asterabad) in silvis prope Sharbat inter Gorgan etZiarat sd Gauba (W 6823)

170

J KU

B

ERA

ET AL



2006

152

169ndash195

Cardamine impatiens

L is a diploid species with awide distribution in Eurasia (Jalas amp Suominen1994) reported as introduced also from North America(Seymour 1969 Rollins 1993) and southern Africa(Dreyer amp Jordaan 2000) Although several infraspe-cific taxa have been described in the past within thisspecies most of them appear currently only in synon-ymy The sole exception is the taxon originallydescribed as

C pectinata

Pall ex DC by Candolle(1821 264) based on the material collected by Pallasin Persia (today Iran) Apart from Iran (Hedge 1968)it is also reported from the Caucasus Turkey and theBalkan Peninsula (Stoyanov amp Stefanov 1948 Gross-geim 1950 Cullen 1965 Assenov 1970 Ketskhoveli1979 Kit Tan 2002) The taxonomic status of thistaxon however has remained controversial It hasbeen treated either as a separate species (by Gross-geim 1950 Assenov 1970 Ketskhoveli 1979) or as asubspecies (by Stoyanov amp Stefanov 1948 Jones ampAkeroyd 1993) or variety (by Trautvetter 1876Zohary Heyn amp Heller 1980) of

C impatiens

Cullen(1965 443) treating this taxon at the varietal rankargued that lsquointermediates between the two varieties[var

impatiens

and var

pectinata

] are of frequentoccurrencersquo This opinion has been shared also by KitTan (2002 184) and Khatri (1988) although the latterauthor treated

C pectinata

at the species level Otherauthors (eg Assenov 1970) have no doubt that thisshould be a clearly separated species

Although

C impatiens

exhibits a wide distributionacross Eurasia the area of the present occurrence of

C pectinata

is regarded to be restricted to the BalkanPeninsula Turkey Iran and lower altitudes of theCaucasus (see references above) These areas areknown to harbour considerable taxonomic and geneticdiversity that can be attributed to processes associ-ated with Pleistocene glaciations Palaeobotanical andphylogeographical studies have shown that the south-ern peninsulas of Iberia Apennines Balkans and theCaspianCaucasus region served as refugia in whichmany European temperate species survived unfavour-able climatic conditions during the glacial periods(Hewitt 2004) Repeated cycles of area contractionand expansion as well as altitudinal shifts in south-ern mountain ranges allowed considerable geneticdiversity to be retained in these regions in contrastwith the genetic impoverishment often observed inmore northern areas (Hewitt 2002 2004) Severalcontinental-scale phylogeographies documentingthese processes have been published in recent yearsaddressing mainly tree species (eg King amp Ferris1998 Fineschi

et al

2002 Hampe

et al

2003Heuertz

et al

2004) A few phylogeographical studiesalso covered or at least marginally included theeasternmost putative refugial areas in Turkey andthe CaspianCaucasus region (King amp Ferris 1998

Hampe

et al

2003 Marcussen 2003 Albach Mar-tiacutenez-Ortega amp Chase 2004) Reconstructions of thevegetation in the last glacial maximum have docu-mented the occurrence of broad-leaved deciduous andmixed forests in the area along the Black Sea coast ofTurkey and Georgia (Adams 1997) This region char-acterized by many Tertiary and Quaternary relic spe-cies (Denk Frotzler amp Davitashvilli 2001) may haveprovided favourable habitats for the survival of sev-eral temperate herbaceous species including the

Car-damine

taxa studied hereTwo main questions are addressed in this study

First we aimed to resolve the taxonomic position ofpopulations assigned to

C pectinata

in respect of thewidespread

C impatiens

Detailed morphometric andmolecular (amplified fragment length polymorphismAFLP) analyses have been carried out together withthe survey of herbarium records from the sympatricregions Second as an introduction to the phylogeog-raphy of both species the distribution of genetic diver-sity in the studied populations and geographicalregions was assessed using AFLP data

MATERIAL AND METHODS

P

LANT

MATERIAL

Population sampling was focused on the area with thereported occurrence of both taxa

C impatiens

and

C pectinata

ie the Balkan Peninsula northern Tur-key and the Caucasus region

C impatiens

was alsosampled from numerous populations in more distantparts of its distribution range (central and southernEurope Japan) to cover the area without the occur-rence of

C pectinata

(Table 1 Fig 1) For morphomet-ric analyses herbarium specimens of a total of 565individuals originating from 26 populations were used(6ndash34 plants per population) These were collected inthe sympatric area of the two taxa Material for AFLPanalyses (silica gel-dried leaves) was represented by82 samples from 24 localities also sampled for mor-phometric studies and 65 samples from an additional23 populations collected mainly in more remote geo-graphical areas (Table 1) For chromosome countsseeds originating from 15 localities were used Thesewere either collected directly in the field or fromplants collected at the localities and transplanted inthe experimental field in Bratislava Slovak Republic

Details on the origins of the plant material are givenin Table 1 Voucher specimens are deposited in theherbarium SAV

To present a detailed view on the distribution ofplants corresponding to

C impatiens

and

C pectinata

especially in the area of their sympatric occurrenceherbarium material from the following herbaria wasrevised B BM G GH K KW LE LEU P PR PRC

TAXONOMY OF

C IMPATIENS

AND

C PECTINATA

171



2006

152

169ndash195

Table 1


Cardamine impatiens

and

C pectinata


n


C impatiens


HK

HKato

HKu

H Kudoh

JK

J Ku

d

era

JL

J Lihovaacute

KM

K Marhold

MK

M Kolniacutek

MO

M Onette

MS

M Slovaacutek

SK

S Kato

SS

S Stoyanov

VGO

V G Onipchenko


n

Morphno ind

AFLP no ind

Cardamine impatiens


42

deg

32

prime

35

Prime

N 23

deg

29

prime

02

Prime

E 660 m 1vi2003

JK amp SS

16 30 1



ndash 30 3


ndash 7 3


ndash 24 3


ndash 30 1


ndash 8 2


16 29 3


16 25 3


16 12 3


ndash ndash 3


16 ndash 3


ndash ndash 3


ndash ndash 3


ndash 21 5


ndash 10 4


ndash ndash 3


16 ndash 3


ndash 22 ndash


ndash 6 3


ndash 8 ndash



g

g

t

172 J KUBERA ET AL





ndash ndash 2


ndash ndash 3


ndash ndash 3


ndash ndash 2


ndash ndash 3


ndash ndash 3


ndash ndash 3


ndash ndash 3


ndash ndash 1


ndash ndash 3


16 ndash ndash


16 ndash ndash




16 30 5


16 34 5


ndash 30 3


ndash 30 1


16 13 2


ndash 25 5


16 28 4


16 29 5


ndash 29 3


ndash 7 5


16 ndash 4


ndash 18 7


AFLP no ind

g

g

g

g

g

g

Table 1 Continued




A

B

174 J KUBERA ET AL
















AFLP FINGERPRINTING



AFLP DATA ANALYSES






176 J KUBERA ET AL



Dis

sim

ilarit

y

6

5

4

3

2

1

0

I-Bu-

1

I-Bu-

3

I-Bu-

4

I-Ro-

20

I-Bu-

2

I-Ro-

18

I-Ge-

15

I-Bu-

5

I-Bu-

8

I-Ro-

19

I-Bu-

7

I-Ge-

14

I-Bu-

6

I-Bu-

9

P-B

u-1

P-B

u-2

P-B

u-4

P-B

u-6

P-B

u-5

P-B

u-3

P-G

e-13

P-T

u-10

P-T

u-11

P-T

u-9

P-T

u-8

P-T

u-7

C impatiens

C pectinata


RESULTS

CHROMOSOME NUMBERS









AFLP ANALYSES






-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






TAXONOMY OF

C IMPATIENS

AND

C PECTINATA

171



2006

152

169ndash195

Table 1


Cardamine impatiens

and

C pectinata


n


C impatiens


HK

HKato

HKu

H Kudoh

JK

J Ku

d

era

JL

J Lihovaacute

KM

K Marhold

MK

M Kolniacutek

MO

M Onette

MS

M Slovaacutek

SK

S Kato

SS

S Stoyanov

VGO

V G Onipchenko


n

Morphno ind

AFLP no ind

Cardamine impatiens


42

deg

32

prime

35

Prime

N 23

deg

29

prime

02

Prime

E 660 m 1vi2003

JK amp SS

16 30 1



ndash 30 3


ndash 7 3


ndash 24 3


ndash 30 1


ndash 8 2


16 29 3


16 25 3


16 12 3


ndash ndash 3


16 ndash 3


ndash ndash 3


ndash ndash 3


ndash 21 5


ndash 10 4


ndash ndash 3


16 ndash 3


ndash 22 ndash


ndash 6 3


ndash 8 ndash



g

g

t

172 J KUBERA ET AL





ndash ndash 2


ndash ndash 3


ndash ndash 3


ndash ndash 2


ndash ndash 3


ndash ndash 3


ndash ndash 3


ndash ndash 3


ndash ndash 1


ndash ndash 3


16 ndash ndash


16 ndash ndash




16 30 5


16 34 5


ndash 30 3


ndash 30 1


16 13 2


ndash 25 5


16 28 4


16 29 5


ndash 29 3


ndash 7 5


16 ndash 4


ndash 18 7


AFLP no ind

g

g

g

g

g

g

Table 1 Continued




A

B

174 J KUBERA ET AL
















AFLP FINGERPRINTING



AFLP DATA ANALYSES






176 J KUBERA ET AL



Dis

sim

ilarit

y

6

5

4

3

2

1

0

I-Bu-

1

I-Bu-

3

I-Bu-

4

I-Ro-

20

I-Bu-

2

I-Ro-

18

I-Ge-

15

I-Bu-

5

I-Bu-

8

I-Ro-

19

I-Bu-

7

I-Ge-

14

I-Bu-

6

I-Bu-

9

P-B

u-1

P-B

u-2

P-B

u-4

P-B

u-6

P-B

u-5

P-B

u-3

P-G

e-13

P-T

u-10

P-T

u-11

P-T

u-9

P-T

u-8

P-T

u-7

C impatiens

C pectinata


RESULTS

CHROMOSOME NUMBERS









AFLP ANALYSES






-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






172 J KUBERA ET AL





ndash ndash 2


ndash ndash 3


ndash ndash 3


ndash ndash 2


ndash ndash 3


ndash ndash 3


ndash ndash 3


ndash ndash 3


ndash ndash 1


ndash ndash 3


16 ndash ndash


16 ndash ndash




16 30 5


16 34 5


ndash 30 3


ndash 30 1


16 13 2


ndash 25 5


16 28 4


16 29 5


ndash 29 3


ndash 7 5


16 ndash 4


ndash 18 7


AFLP no ind

g

g

g

g

g

g

Table 1 Continued




A

B

174 J KUBERA ET AL
















AFLP FINGERPRINTING



AFLP DATA ANALYSES






176 J KUBERA ET AL



Dis

sim

ilarit

y

6

5

4

3

2

1

0

I-Bu-

1

I-Bu-

3

I-Bu-

4

I-Ro-

20

I-Bu-

2

I-Ro-

18

I-Ge-

15

I-Bu-

5

I-Bu-

8

I-Ro-

19

I-Bu-

7

I-Ge-

14

I-Bu-

6

I-Bu-

9

P-B

u-1

P-B

u-2

P-B

u-4

P-B

u-6

P-B

u-5

P-B

u-3

P-G

e-13

P-T

u-10

P-T

u-11

P-T

u-9

P-T

u-8

P-T

u-7

C impatiens

C pectinata


RESULTS

CHROMOSOME NUMBERS









AFLP ANALYSES






-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e









A

B

174 J KUBERA ET AL
















AFLP FINGERPRINTING



AFLP DATA ANALYSES






176 J KUBERA ET AL



Dis

sim

ilarit

y

6

5

4

3

2

1

0

I-Bu-

1

I-Bu-

3

I-Bu-

4

I-Ro-

20

I-Bu-

2

I-Ro-

18

I-Ge-

15

I-Bu-

5

I-Bu-

8

I-Ro-

19

I-Bu-

7

I-Ge-

14

I-Bu-

6

I-Bu-

9

P-B

u-1

P-B

u-2

P-B

u-4

P-B

u-6

P-B

u-5

P-B

u-3

P-G

e-13

P-T

u-10

P-T

u-11

P-T

u-9

P-T

u-8

P-T

u-7

C impatiens

C pectinata


RESULTS

CHROMOSOME NUMBERS









AFLP ANALYSES






-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






174 J KUBERA ET AL
















AFLP FINGERPRINTING



AFLP DATA ANALYSES






176 J KUBERA ET AL



Dis

sim

ilarit

y

6

5

4

3

2

1

0

I-Bu-

1

I-Bu-

3

I-Bu-

4

I-Ro-

20

I-Bu-

2

I-Ro-

18

I-Ge-

15

I-Bu-

5

I-Bu-

8

I-Ro-

19

I-Bu-

7

I-Ge-

14

I-Bu-

6

I-Bu-

9

P-B

u-1

P-B

u-2

P-B

u-4

P-B

u-6

P-B

u-5

P-B

u-3

P-G

e-13

P-T

u-10

P-T

u-11

P-T

u-9

P-T

u-8

P-T

u-7

C impatiens

C pectinata


RESULTS

CHROMOSOME NUMBERS









AFLP ANALYSES






-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e










AFLP FINGERPRINTING



AFLP DATA ANALYSES






176 J KUBERA ET AL



Dis

sim

ilarit

y

6

5

4

3

2

1

0

I-Bu-

1

I-Bu-

3

I-Bu-

4

I-Ro-

20

I-Bu-

2

I-Ro-

18

I-Ge-

15

I-Bu-

5

I-Bu-

8

I-Ro-

19

I-Bu-

7

I-Ge-

14

I-Bu-

6

I-Bu-

9

P-B

u-1

P-B

u-2

P-B

u-4

P-B

u-6

P-B

u-5

P-B

u-3

P-G

e-13

P-T

u-10

P-T

u-11

P-T

u-9

P-T

u-8

P-T

u-7

C impatiens

C pectinata


RESULTS

CHROMOSOME NUMBERS









AFLP ANALYSES






-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






176 J KUBERA ET AL



Dis

sim

ilarit

y

6

5

4

3

2

1

0

I-Bu-

1

I-Bu-

3

I-Bu-

4

I-Ro-

20

I-Bu-

2

I-Ro-

18

I-Ge-

15

I-Bu-

5

I-Bu-

8

I-Ro-

19

I-Bu-

7

I-Ge-

14

I-Bu-

6

I-Bu-

9

P-B

u-1

P-B

u-2

P-B

u-4

P-B

u-6

P-B

u-5

P-B

u-3

P-G

e-13

P-T

u-10

P-T

u-11

P-T

u-9

P-T

u-8

P-T

u-7

C impatiens

C pectinata


RESULTS

CHROMOSOME NUMBERS









AFLP ANALYSES






-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e










AFLP ANALYSES






-447

-053

340

734

Prin2

456

130

-195

-521

Prin1

-401

-128

145

419

Prin3

178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






178 J KUBERA ET AL





DISCUSSION




FREQUENCY

0

10

20

30

40

50

60

70

80

90

100

110

120

130

140

150

Can1

-66 -54 -42 -30 -18 -06 06 18 30 42 54 66




0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e









0

10152025303540455055606570758085

5

heig

ht o

f st

em (

cm)

5

10

15

20

25

30

num

ber

of s

tem

leav

es

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

num

ber

of le

afle

ts o

f th

e m

iddl

e st

em le

af

15

20

25

30

35

40

45

50

55

60

65

70

leng

th o

f pe

tals

(m

m)

04

07

10

13

16

19

22

25

28

wid

th o

f pe

tals

(m

m)

08

10

12

14

16

18

20

22

24

26

28

30

32

leng

th o

f se

pals

(m

m)

10

15

20

25

30

35

40

45

leng

th o

f fil

amen

ts (

mm

)

0

5

10

15

20

25

30

35

40

45

50

55

LLL

WLL

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a

impa

tiens

pect

inat

a




180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






180 J KUBERA ET AL



I - It - 27I - It - 27

I - It - 28I - Sp - 30

I - Sp - 30I - It - 28I - Cr - 24I - Cr - 24

I - Cr - 24I - It - 26

I - It - 26I - It - 26I - It - 26

I - Sk - 33I - Sk - 33

I - Sk - 33I - Sw - 29I - Sw - 29

I - Sw - 29

I - Bu - 5I - It - 26

I - It - 28I - Sk - 32I - Sk - 32I - Sk - 32

I - Bu - 11I - Bu - 11

I - Bu - 11I - Bu - 4

I - Bu - 4I - Bu - 4

I - Bu - 3

I - Bu - 3I - Bu - 6

I - Bu - 6I - Bu - 7I - Bu - 7

I - Bu - 7I - Bu - 2I - Ro - 22

I - Ro - 22I - Sk - 34I - Sk - 34

I - Sk - 34I - Bu - 9I - Bu - 9I - Bu - 9

I - Se - 25I - Se - 25

I - Se - 25I - Bu - 9

I - Bu - 9I - Bu - 9

I - Bu - 2I - Bu - 2

I - Bu - 10I - Bu - 10I - Bu - 10

I - Bu - 3I - Ro - 21

I - Ro - 21I - Ro - 21

I - Ro - 23I - Sk - 31

I - Sk - 31I - Sk - 31

I - Bu - 1I - Ro - 19I - Ro - 19

I - Ro - 19I - Ru - 16

I - Ru - 16I - Ru - 16

I - Tu - 12I - Tu - 12

I - Tu - 12I - Ge - 15

I - Ge - 15I - Ge - 15I - Ge - 15

I - Ge - 14I - Ge - 14I - Ge - 14

I - Ge - 14I - Ge - 14

I - Ru - 17I - Ru - 17

I - Tu -13I - Tu -13

I - Tu -13I - Ru - 17

I - Ja - 35I - Ja - 36I - Ja - 36

I - Ja - 36P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Ge - 13P - Ge - 13

P - Tu - 9P - Tu - 9P - Tu - 9

P - Tu - 9

P - Tu - 9

P - Tu -8P - Tu -8

P - Tu -8

P - Tu -8P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 7P - Tu - 7

P - Tu - 11P - Tu - 11P - Tu - 11P - Tu - 11

P - Tu - 11

P - Bu - 4

P - Bu - 4

P - Bu - 4

P - Bu - 2

P - Bu - 2

P - Bu - 2

P - Bu - 2P - Bu - 2

P - Bu - 3P - Bu - 3

P - Bu - 3P - Bu - 3

P - Bu - 3

P - Bu - 6

P - Bu - 6P - Bu - 1

P - Bu - 1

P - Bu - 1P - Bu - 5

P - Tu - 12P - Tu - 12P - Tu - 12

P - Tu - 12P - Tu - 10

P - Tu - 10P - Tu - 10

C pectinata

C impatiens

9595

86

9570

6453

8299

91 66

59

61

62

81

82

7370

77

93

57

80

6889

7865

81

91

74 79

5799

6410076

61

52

54

54

76

6874

76

100

100

55

72

73

100

lazicalazica

lazica

0005 changes





Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e










Variancecomponents

of totalvariance
















182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






182 J KUBERA ET AL






















++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e













++++++P-Tu-10

P-Tu-7

P-Tu-8

P-Tu-9

+++++

P-Tu-11++

P-Ge-13+++

P-Tu-12+++++

P-Bu-1

++++

P-Bu-2+

P-Bu-5P-Bu-6

P-Bu-3

P-Bu-4


western group

I-Sp-30

A

B

I-It-27

I-It-28

I-Sw-29

I-It-26

+++

I-Sk-33

I-Sk-31++++

I-Sk-34I-Sk-32

+

low HSh

medium HSh

high HSh


I-Cr-24I-Se-25

+

++I-Ro-22

I-Ro-23

I-Ro-21I-Ro-19

I-Bu-11

I-Bu-2

I-Bu-6

I-Bu-789

I-Bu-3410+ +

+

++++I-Tu-12I-Tu-13

I-Ru-16

I-Ru-17 +

I-Ge-14

I-Ge-15+


184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






184 J KUBERA ET AL






g

















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e





















186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






186 J KUBERA ET AL































ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e

















ACKNOWLEDGEMENTS


REFERENCES











188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






188 J KUBERA ET AL










































































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e











































190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






190 J KUBERA ET AL


APPENDIX







g

g

g

e

e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e











192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e






192 J KUBERA ET AL











r

g

g



194 J KUBERA ET AL



e









r

g

g



194 J KUBERA ET AL



e






194 J KUBERA ET AL



e










Documents

Taxonomy and phylogeography of Cardamine impatiens …ibot.sav.sk/usr/Karol/docs/PDF_files/Kucera et al. BJLS_2006.pdf · Taxonomy and phylogeography of Cardamine impatiens ... did