Molecular phylogenetics, morphology and a revised classification of the complex genus Saxifraga (Saxifragaceae)

Abstract

Saxifraga, the most species-rich and taxonomically complex genus of Saxifragaceae, is a characteristic component of temperate to polar climatic zones and of montane to alpine vegetation belts in mountain ranges of the Northern Hemisphere. The genus encompasses more than 440 species, which display notable diversity in growth form, vegetative and reproductive characters, as well as in micromorphology of pollen grains and seeds. Using a large taxon set including representatives attributed to nearly all recognised subgenera, sections and subsections of Saxifraga (altogether 254 species), as well as a broad coverage of outgroup taxa, we inferred evolutionary relationships within Saxifraga, explored the reasons of the striking inconsistencies between previous taxonomic treatments of Saxifraga, estimated the degree of homoplasy in characters frequently employed for classification, assessed the role of hybridisation in the evolution of the genus and, finally, provided a phylogeny-based framework upon which to revise its taxonomy. The phylogenetic trees based on nuclear ribosomal internal transcribed spacer (ITS) and plastid trnL-trnF DNA markers, including more than 460 newly generated sequences, were generally highly congruent, except within a single clade (sect. Saxifraga), in which incomplete lineage sorting and presumably allopolyploid speciation plays an important evolutionary role. Major lineages of Saxifraga are concordantly resolved by the DNA markers from both organelles with mostly strong node support. Our molecular phylogenetic results support the recognition of at least 13 sections and 9 subsections within Saxifraga. A part of these lineages agrees well with previously recognised infrageneric groupings, whereas others are differently delineated. Some of the groups identified by molecular phylogenetics are characterised only by the combination of different morphological characters. Frequently, micromorphological characters support the clades in the phylogenetic tree comparatively well. We here provide new ranks for three infrageneric epithets, namely for sect. Saxifraga subsect. Androsaceae and subsect. Arachnoideae and sect. Porphyrion subsect. Squarrosae. Finally, our study identifies remaining taxonomic uncertainties within Saxifraga.

Full text

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Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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
 || 
INTRODUCTION
   Saxifraga

 
 
    
   


Artemisia Carex  RanunculusPoa
 
 S. hederifo-
lia S. gemmipara 
       S. adenodes
   S. magellanica  S. boussingaultii
 S. pavonii S. bicuspidata

 Saxifraga
 S Micranthes 
Molecular phylogenetics, morphology and a revised classification of
the complex genus Saxifraga (Saxifragaceae)
Natalia Tkach,1 Martin Röser,1 Georg Miehe,2 Alexandra N. Muellner-Riehl,3,4 Jana Ebersbach,3
Adrien Favre3 & Matthias H. Hoffmann1
 Martin Luther Universit y Halle-Wittenberg, Institute of Biology, Geobotany and Botanical Garden, Neuwerk 21, 06108 Halle,
Germany
 Philipps-University Marburg, Department of Geography, Deutschhausstraße 10, 35032 Marburg, Germany
 University of Leipzig, Institute of Biology, Department of Molecular Evolution and Plant Systematics, Herbarium LZ, Johannisallee
21–23, 04103 Leipzig, Germany
 German Centre for Integrative Biodiversit y Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, 04103 Leipzig, Germany
 Natalia Tkach, natalia.tkach@botanik.uni-halle.de; Martin Röser, martin.roeser@botanik.uni-halle.de;
Matthias H. Hoffmann, matthias.hoffmann@botanik.uni-halle.de
DOI
 
Abstract
Saxifraga



Saxifraga
Saxifraga
Saxifraga


trnL-trnF
Saxifraga
Sa xifraga

Saxifraga
 

Sa xifraga
Androsaceae ArachnoideaePorphyrionSquarrosae  
Saxifraga.
Keywords

Supplementary Material



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–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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Micranthes

Micranthes
Chrysosplenium
Boykinia  Heuchera
Mitella  
Saxifraga
   
  

 
Saxifraga

Saxifraga
. Saxifraga Ciliatae

LigulataeTrachyphyl-
lum 
    
Bronchiales 

S. rivularis
  
S. oppositifolia





 
Saxifraga
  
Saxifraga



Saxifraga    
  




Bronchiales


    
  Saxifraga

Saxifraga





Saxifraga   
  
  
    


    
 S. cernua 
   Saxifraga   
 S. oppositifolia
 S. mutataS. aizoides 
 
nS. eschscholtziin
S. androsacea
 

Saxifraga




Saxifraga




Saxifraga 



Saxifraga 
  

 Saxifraga

 


 

S. bicuspidataS. korshinskyi S. mertensiana
 S. sieversiana Cascadia nuttallii 




Saxi-
fraga.

MATERIALS AND METHODS
Sampling strategy, plant material and taxonomic treat-
ments. —
 Saxifraga
      
 

   


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Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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–
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Fig. 1.
Saxifraga.
A,
S. veitchiana Irregulares
B,
S. aristulata      Ciliatae    
C,
S. filicaulis     Ciliatae   

D–E,
S. signataCiliatae
F,
S. diversifoliaCiliatae

G,
S. serpyllifoliaCiliatae
H,
S. umbrosaGymnopera
M. Röser 10581 
I,
S. bryoidesTrachyphyllum 
J,
S. cotyledon L. 
LigulataeM. Röser 10387


1162
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–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record

1163
Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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–
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Odontophyllae S. odontophylla
Tetrameridium


 


Saxifraga

       

     
 
Pterostemon rotundifoliusItea virg-
inica
 
 



  


 









Laboratory work and DNA sequence handling. —
 

trnL-trnF
trnL trnLtrnF
     

Saxifraga





  

trnL-trnF
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L
trnL
trnF
   
  
    



    



trnL-
trnF 
 




 






  

 
 




 



Fig. 2.
Saxifraga.
A,
S. caesiaPorphyrion Squarrosae
B,
S. juniperi-
foliaPorphyrionKabschiaM. Röser 2779
C,
S. aizoides L.
 Porphyrion Mutatae   M. Röser 2182 
D,
S. oppositifolia 
rudolphianaPorphyrion Oppositifoliae

E,
S. biflora Porphyrion Oppositifoliae
F,
S. mutataPorphy-
rionMutataeM. Röser 10412
G,
S. adscendensSaxifragaTridactylites
 M. Röser 9825 
H,
S. androsaceaSaxifraga Androsaceae

I,
S. sedoideshohenwartii  SaxifragaArachnoideae
M. Röser 10442 
J,
S. arachnoidea SaxifragaArachnoideae
M. Röser 10259
K,
S. bourgaeanaSaxifragaSaxifragaM. Röser
4682
L,
S. geranioidesSaxifragaSaxifragaM. Röser
10531 
M,
S. pubescens  iratianaSaxifragaSaxifraga 


1164
TAX ON
–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record
Table 1.
Saxifraga





 
 Irregulares
S. stolonifera 
sect. Diptera
S. stolonifera
Boraphila 
Punctatae
sect. Irregulares
S. stolonifera
sect. Heterisia

S. mertensiana 
subg. Diptera
S. stolonifera
sect. IrregularesS. stolonifera
sect. HeterisiaS. mertensi-
ana
Saxifragella
S. bicuspi-
data
Saxifragella bicuspidata

Saxifragella bicuspidata Saxifragella bicuspidata
 Pseudocymbalaria
S. sieversiana 
Dactyloides 
Eudactyloideae

Cespitosae 

Boraphila
Punctatae
Saxifraga
sect. Pseudocymbalaria
S. sieversiana
sect. Komarovia
S. korshinskyi
Tridactylites 

subsect. Lacteae
S. lactea 
 Bronchiales
S. bronchialis
Trachyphyllum 

Trachyphyllum Porphyrion

Trachyphyllum
subsect. Xeromorphicae
S. bronchialis
subsect. Criomorphicae
S. funstonii

subsect. Dentaphyllae

S. tricuspidata 
 Ciliatae
S. hirculus
sect. Hirculus
S. hirculus
grex Flagellares
S. flagellaris

grex Gemmiparae

S. gemmipara
grex Densifoliatae 
S. macrostigma

grex Hemisphaericae S. hemi-
sphaerica
grex Hirculoideae 

S. nigroglandulosa
grex Lychnitidae 

S. lychnitis
grex Nutantes 

S. nutans
grex Sediformis 

S. sediformis
grex Turfosae 

S. turfosa
sect. CiliataeS. hirculus
subsect. Flagellares

S. flagellaris
subsect. Gemmiparae 
S. gemmipara
subsect. Hemisphaericae 
S. hemisphaerica

subsect. Hirculoideae
S. nigroglandulosa 

subsect. Rosulares 
S. sediformis
subsect. Serpyllifoliae
S. serpyllifoliae 
subg. Hirculus
S. hirculus
sect. Aretiaria
S. eschscholtzii
sect. CiliataeS. hirculus
sect. Flagellares 
S. flagellaris
sect. Rosulares
S. sediformis


1165
Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record
Tab le 1.





 
 Cymbalaria
S. cymbalaria 
sect. CymbalariaS. cymbalariasect. CymbalariaS. cymbalaria Saxifraga
sect. Cymbalaria
S. cymbalaria
 Cotylea
S. rotundifolia
sect. Miscopetalum
S. rotundifolia
sect. CotyleaS. rotundifolia Saxifraga
sect. CotyleaS. rotundifolia
 Gymnopera
S. hirsuta
sect. Robertsonia
S. hirsuta
sect. GymnoperaS. hirsuta Porphyrion
sect. Gymnopera
 Mesogyne
S. sibirica
Nephrophyllum

grex Sibiricae
S. sibirica
sect. MesogyneS. sibirica Saxifraga
Saxifraga
subsect. Mesogyne
S. sibirica
 Trachyphyllum
S. aspera
TrachyphyllumS. aspera TrachyphyllumS. aspera Porphyrion
TrachyphyllumS. aspera
subsect. Fibrosophyllae

S. aspera
 Ligulatae
S. cotyledon
Euaizonia
S. paniculata
grex Cotyledoniae
S. cotyledon
grex Peraizoonia
S. paniculata
grex Crustatae
S. crustata
LigulataeS. cotyledon
subsect. Aizoonia
S. paniculata
Porphyrion
LigulataeS. cotyledon
 Porphyrion
S. oppositifolia
sect. Porphyrion
S. oppositifolia
sect. Xanthizoon
S. aizoides
sect. KabschiaS. marginata

Euaizoonia 
grex Florulentae
S. florulenta
grex Mutatae
S. mutata
sect. PorphyrionS. oppositifolia
sect. XanthizoonS. aizoides
Ligulatae
subsect. Florulentae
S. florulenta

subsect. Mutatae
S. mutata
PorphyrionS. oppositifolia
sect. PorphyrionS. oppositi-
folia
sect. XanthizoonS. aizoides
sect. KabschiaS. marginata
Ligulatae
Squarrosae 

.
S. squarrosa
Kabschia
grex Squarrosae
S. squarrosa
Porphyrion
Kabschia

ser. Squarrosae
S. squar-
rosa
Porphyrion
Kabschia
Mutatae
S. mutata
sect. Xanthizoon
S. aizoides
Euaizoonia 
grex MutataeS. mutata
sect. XanthizoonS. aizoides
Ligulatae
subsect. MutataeS. mutata
Porphyrion
sect. XanthizoonS. aizoides
Ligulatae
Oppositifoliae
S. oppositifolia
sect. PorphyrionS. oppositifolia Porphyrion
Oppositifoliae
S. oppositifolia
Porphyrion
sect. PorphyrionS. oppositi-
folia
 Florulentae 
S. florulenta

Euaizoonia
grex FlorulentaeS. floru-
lenta
Ligulatae
subsect. Florulentae
S. florulenta
Porphyrion
Ligulatae

1166
TAX ON
–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record
Tab le 1.





 
 Kabschia
S. marginata 
KabschiaS. marginata
grex Mediae
S. media
grex Juniperifoliae
S. juniperifolia
grex Kotschyanae
S. kotschyi
grex Marginatae
S. marginata
grex Rigidae
S. vandellii
grex Aretitoidae
S. aretitoides
Porphyrion
subsect. Engleria
S. media
KabschiaS. marginata
ser. Juniperifoliae
S. juniperi-
folia
ser. Lilacinae
S. lilacina

ser. Marginatae
S. mar-
ginata
ser. Rigidae
S. vandellii
ser. Aretitoideae
S. aretitoi-
des
Porphyrion
Kabschia S. marginata
 Saxifraga S. granulata TridactylitesS. tri dac-
tylites
DactyloidesS. granulata
Holophyllae

grex Tenellae
S. tenella
grex Sedoideae
S. sedoides 
grex Aphyllae
S. aphylla
grex Androsaceae
S. androsacea
Eudactyloideae
grex Axilliflorae
S. praeter-
missa
grex Aquaticae
S. aquatica
.
grex Ceratophyllae
S. trifurcata

grex Gemmiferae
S. globu-
lifera
CespitosaeS. cespitosa
grex Exarato-Moschatae
S. exarata

NephrophyllumS. granulata
grex Arachnoideae
S. arachnoidea
grex Irriguae
S. irrigua
grex Granulatae
S. granulata
sect. SaxifragaS. granulata
subsect. Tridactylites
S. tridactylites
subsect. SaxifragaS. granu-
lata
subsect. Holophyllae
subsect. Triplinervium
S. cespitosa 
SaxifragaS. granulata
TridactylitesS. tridac-
tylites
subsect. Tridactylites
S. tridactylites
sect. Discogyne
S. paradoxa
sect. Stoloniferae
S. geranioides
SaxifragaS. granulata
subsect. Saxifraga
S. granulata
Tridactylites 
S. tridactylites
Dactyloides
Eudactyloideae
Axilliflorae
TridactylitesS. tridactylites
Saxifraga
Triplinervium 

subsect. TridactylitesS. tridac-
tylites
Saxifraga
TridactylitesS. tridac-
tylites
subsect. TridactylitesS. tri-
dactylites


Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record
Tab le 1.





 
Androsaceae 

.
S. androsacea
Dactyloides
Holophyllae
grex AndrosaceaeS. andro-
sacea
Saxifraga
Holophyllae
Triplinervium
Saxifraga
Discogyne
Stoloniferae
Arachnoideae 


S. arachnoidea
Tridactylites
Dactyloides
Holophyllae
grex TenellaeS. tenella
grex SedoideaeS. sedoi-
des
grex AphyllaeS. aphylla
Nephrophyllum
grex Arachnoideae
S. arachnoidea
Saxifraga
Triplinervium
Holophyllae
Saxifraga
DiscogyneS. paradoxa
Stoloniferae
Saxifraga S. granu-
lata
DactyloidesS. granulata
Eudactyloideae
AxillifloraeS. praeter-
missa
grex AquaticaeS. aquatica
grex Ceratophyllae
S. trifurcata
grex GemmiferaeS. globu-
lifera
CespitosaeS. cespitosa
grex Exarato-Moschatae
S. exarata
Nephyrophyllum
Irriguae
grex GranulataeS. granulata
SaxifragaS. granulata
subsect. SaxifragaS. granu-
lata
Triplinervium
S. cespitosa
SaxifragaS. granulata
SaxifragaS. granulata
StoloniferaeS. gerani-
oides
Cascadia nuttallii

Tridactylites Cascadia nuttallii Cascadia nuttallii
Incertae sedis
S. odontophylla
S. nana 
Boraphila
Punctatae
Tetrameridium
S. nana
Odontophyllae
Porphyrion
Oppositifoliae
Saxifraga
Odontophyllae



 
  S. mertensiana.
 S. sieversianaS. selemdzhensis
 S. lactea.
  S. sieversianaS. korshinskyi.
 Stellariifoliae Cinerascentes

 Cinerascentes
 Cinerascentes
 
 OppositifoliaeTetrameridiumTetrameridium
Oppositifoliae 
 S. nuttalliiCascadia nuttallii.
 MuscoideaeGlabellae
13 Saxifraga muscoides
 S. irrigua 


TAX ON
–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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Phylogenetic analyses. —




 
   



  

  
  
  






 


Saxifraga 

 
     

  
   




RESU LT S
Alignment characteristics and phylogenetic analyses. —

SaxifragatrnL-trnF
Saxi-
fraga

Saxifraga
trnL-trnF
Saxifraga
 
         

  

Saxifraga




L =  L
= trnL-trnF

 trnL-trnF 


  
  
   
L = 
   



Tree support and incongruences. —


trnL-trnF

 Saxifraga
 


   




   


[
      

  Saxifraga   

trnL-trnF
 



trnL-trnF
S. wahlenbergii
   
trnL-trnF
S. aphylla

trnL-trnFS. moschata
   
 Saxifraga 




Saxifraga
 
S. cymbalariaS. hederacea S. sib-
thorpii   


 


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Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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–
Version of Record
Phylogenetic relationships between major clades. —

 

Cascadia nuttallii S. nut-
tallii Saxifraga
 

Saxifraga   
   Irregulares
S. mertensianaHeteri-
sia 
Saxifraga
bicuspidata 
  
Bronchiales 
S. sieversianaS. korshinskyi
S. lacteaPseudocymbalaria

Saxifraga  
trnL-trnFPseudocymbalaria
Bronchiales



Ciliatae
Fig. 3.
SaxifragatrnL-
trnF                
SaxifragaCascadia nuttalliiSaxifragodes albowiana
Pterostemon rotundifoliusItea virginica

Nuclear ribosomal ITS
DNA sequence data
Heucheroids
Cascadia nuttallii Cascadia nuttallii
Saxifragodes
albowiana
Heucheroids
Saxifragodes
albowiana
100/100 100/64
69/-
86/85
88/72
94/83
83/60
68/66
74/-
100/76
100/95
100/99
100/100
62/- 62/65
94/80
99/99
100/99
67/69
89/94
75
84
95/98
98/93
93/91
98/91
87/53
70/70
100
100
100
100
100/100
55/67
52/53
62
72
51/-
99/93
Plastid trnL-trnF
DNA sequence data
OUTGROUP
100/100
100/94
66
86
91/82
53
73
96/87
100/90
57/56
100/95
100/95
99/95
100/100
100/100
100/100
99/100
100
100
86/89
100/97
100/100
100/100
100/100
100/100
100/100
100/100
100/100
100/100
100/100
100/100
100/100
100/100
100/100
50/69
100/100
100/100
100/100
100/100
Combined dataset
ITS and trnL-trnF
Sections of Saxifraga /
main clades
Sections of Saxifraga /
main clades
ML / MP bootstrap values
SAXIFRAGA
PORPHYRION
LIGULATAE
TRACHY-
PHYLLUM
MESOGYNE
GYMNOPERA
COTYLEA
CYMBALARIA
CILIATAE
BRONCHIALES
SAXIFRAGELLA
SAXIFRAGELLA
IRREGULARES
OUTGROUP
SAXIFRAGA
PORPHYRION
LIGULATAE
TRACHY-
PHYLLUM
MESOGYNE
GYMNOPERA
COTYLEA
CYMBALARIA
CILIATAE
BRONCHIALES
PSEUDO-
CYMBALARIA
IRREGULARES
PSEUDO-
CYMBALARIA
SaxifragaSaxifraga

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–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record
 Cymbalaria 
Saxifraga   
trnL-trnFCiliatae
Cymbalaria   



Saxifraga 
Saxifraga 

Saxifraga
Cotylea
Gymnopera Ligulatae Mesogyne
PorphyrionTrachyphyllum




Gymnopera   
trnL-trnF
E 
Porphyrion   

Mesogyne  
Fig. 4A–D.
Saxifraga
 trnL-trnF Pterostemon rotundifolius Itea virginica     
Saxifraga irrigua
Saxifraga

S. cespitosa subsp. monticola
S. cespitosa subsp. cespitosa
S. terektensis
S. rosacea
S. hypnoides
S. continentales
S. cuneata
S. fragosoi
S. maderensis
S. portosancta
S. trifurcata
S. babiana
S. camposii var. leptophylla
S. canaliculata
S. latepetiolata (2)
S. latepetiolata (1)
S. dichotoma
S. carpetana
S. bulbifera
S. granulata
S. moschata
S. graeca
S. pentadactylis subsp. willkommiana
S. pentadactylis (1)
S. pentadactylis (2)
S. nevadensis
S. geranioides
S. magellanica
S. moncayensis
S. genesiana
S. vayredana
S. cebennensis
S. exarata
S. pontica
S. praetermissa
S. pubescens subsp. iratiana
S. intricata
S. hariotii
S. fragilis (2)
S. fragilis (1)
S. aquatica
S. pedemontana subsp. cymosa
S. reuteriana
S. globulifera (1)
S. globulifera (2)
S. erioblasta
S. trabutiana
S. wahlenbergii
S. osloensis (1)
S. tridactylites
S. discolor
S. rigoi
S. corsica
S. bourgaeana
S. cossoniana
S. gemmulosa
S. cintrana
S. tenella (2)
S. sedoides subsp. hohenwartii
S. tenella (1)
S. paradoxa
S. berica
S. presolanensis
S. petraea
S. arachnoidea
S. androsacea
S. seguieri
S. aphylla
S. depressa
S. irrigua
S. adscendens
S. italica
S. osloensis (2)
S. stolitzkae
S. andersonii
S. matta-florida
S. georgei
S. alpigena
S. lilacina
S. pulvinaria
S. afghanica
S. duthiei
S. roylei
S. unguipetala
S. scardica
S. wendelboi
S. marginata (1)
S. albertii
S. ferdinandi-coburgi
S. marginata (2)
S. kotschyi
S. sancta
S. marginata var. bubakii
S. felineri
S. diapensioides
S. aretioides
S. iranica
S. charadzeae
S. subverticillata
S. vandellii
S. tombeanensis
S. caspica
S. dinnikii
S. caucasica
S. colchica
S. ruprechtiana
S. pseudolaevis
S. juniperifolia
S. scleropoda
S. desoulavyi
S. columnaris
S. stribrnyi
S. thessalica
S. sempervivum
S. federici-augusti subsp. grisebachii
S. porophylla
S. spruneri
S. corymbosa
S. luteo-viridis
S. oppositifolia (1)
S. oppositifolia subsp. asiatica (1)
S. oppositifolia subsp. asiatica (2)
S. oppositifolia subsp. oppositifolia
S. oppositifolia subsp. smalliana
S. oppofitifolia (2)
S. retusa subsp. augustana
S. mutata
S. aizoides
S. squarrosa
S. caesia
S. florulenta
S. valdensis (1)
S. cf. cotyledon
S. valdensis (2)
S. cotyledon
S. cochlearis
S. crustata
S. hostii
S. paniculata
S. cartilaginea
S. kolenatiana
S. lingulata
S. callosa
S. bryoides
S. aspera
S. spathularis (1)
S. coriifolia
S. taygetea
S. rotundifolia
S. spathularis (2)
S. umbrosa
S. hirsuta
S. hederacea (2)
S. hederacea (1)
S. cymbalaria (1)
S. cymbalaria (2)
S. sibthorpii
S. saginoides
S. viscidula
S. aristulata
S. montana
S. pseudohirculus
S. peplidifolia
S. litangensis
S. yarlungzangboensis
S. przewalskii
S. hirculoides
S. tangutica
S. sinomontana
S. heleonastes
S. cf. hirculoides
S. tibetica
S. hirculus var. alpina
S. hirculus var. hirculus (1)
S. hirculus var. hirculus (2)
S. insolens
S. diversifolia var. haematophylla
S. egregia
S. hookeri
S. implicans
S. moorcroftiana
S. latifolia
S. lychnitis
S. isophylla
S. zimmermannii
S. parnassifolia
S. bergenioides
S. caveana
S. diversifolia
S. xiaozhongdianensis
S. kingdonii
S. kingiana
S. nanella
S. stella-aurea var. poyadena
S. gemmigera var. gemmuligera
S. contraria f. rubella
S. engleriana
S. unguiculata
S. aurantiaca
S. tatsienluensis
S. perpusilla
S. heterotricha
S. umbellatula var. umbellatula
S. signatella
S. umbellatula var. muricola
S. dingqingensis
S. umbellatula var. pectinata
S. serpyllifolia subsp. glutinosa
S. aleutica
S. serpyllifolia var. serpyllifolia
S. sanguinea
S. chrysantha
S. jaquemontiana
S. finitima
S. punctulata
S. sediformis
S. macrocalyx
S. platysepala
S. setigera
S. flagellaris subsp. crandallii
S. flagellaris subsp. crassiflagellata
S. flagellaris subsp. stenophylla
S. zhidoensis
S. mucronulata
S. mucronulatoides
S. pilifera
S. nigroglandulifera
S. nangxianensis
S. komarovii
S. consanguinea
S. angustata
S. hispidula (1)
S. hispidula (2)
S. strigosa var. ramosa
S. wallichiana
S. gouldii
S. brachypoda
S. gemmipara
S. filicaulis
S. substrigosa
S. eschscholtzii (1)
S. eschscholtzii (2)
S. firma (2)
S. funstonii (2)
S. funstonii (1)
S. nishidae
S. firma (1)
S. verspertina
S. taylori
S. sieversiana
S. korshinskii
S. lactea
S. bicuspidata
S. veitchiana
S. stolonifera
S. rufescens
S. mertensiana
S. nipponica
OUTGROUP
S. fortunei var. incisolobata
S. fortunei
S. cortusifolia
S. omolojensis (2)
S. anadyrensis (2)
S. anadyrensis (1)
S. spinulosa
S. omolojensis (1)
S. kruhsiana
S. stelleriana
S. cherlerioides
S. derbekii
S. bronchialis
S. bronchialis subsp. austromontana
S. caulescens
S. tricuspidata (1)
S. tricuspidata (2)
S. cherlerioides var. rebunshirensis
S. wardii
S. brunonis (1)
S. brunonis (2)
S. cuneifolia
S. blepharophylla
S. biflora
S. rivularis
S. hyperborea
S. radiata (1)
S. radiata (2)
S. bracteata (1)
S. bracteata (2)
S. cernua (2)
S. cernua (1)
S. carpatica
S. sibirica
S. granulifera
S. flexuosa
S. rivularis subsp. arctolitoralis
S. biternata
S. conifera (1)
S. conifera (2)
S. haenseleri
A
B
C
B
91/-
91/90
52/66 56/66
63/-
61/-
99/94
55/65
57/61
50/62
96/ 99
74/64
99/94
95/96
94/97
100/100
100/100
68/64
92/93
74/80
76/76
71/69
100/100
53/-
55/-
100/100
100/100
100/100
100/100
69/-
83/-
61/-
89/94
92/99
100/100
100/99
100/100
100/100
100/100
100/100
100/ 100
100/100
100/98
100/99
75/74
91/ 89
78/ 83
100/100
99/ 100
66/66
71/80
73/-
98/100
97/95
97/-
100/99
81/77
96/92
67/68
76/100
S. cespitosa subsp. monticola +
S. cespitosa subsp. cespitosa +
S. terektensis +
S. rosacea +
S. hypnoides +
S. continentalis +
S. cuneata +
S. fragosoi +
S. maderensis +
S. portosanctana +
S. trifurcata +
S. babiana +
S. camposii var. leptophylla +
S. canaliculata +
S. latepetiolata (2)
S. latepetiolata (1) +
S. dichotoma +
S. carpetana +
S. bulbifera +
S. granulata +
S. moschata +
S. graeca +
S. pentadactylis subsp. willkommiana
S. pentadactylis (1) +
S. pentadactylis (2) +
S. nevadensis +
S. geranioides +
S. magellanica +
S. moncayensis +
S. genesiana +
S. vayredana +
S. cebennensis +
S. exarata +
S. pontica +
S. praetermissa +
S. pubescens subsp. iratiana +
S. intricata
S. hariotii
S. fragilis (2)
S. fragilis (1) +
S. aquatica
S. pedemontana subsp. cymosa +
S. reuteriana
S. globulifera (1) +
S. globulifera (2) +
S. erioblasta +
S. trabutiana +
S. wahlenbergii +
S. osloensis (2) +
S. tridactylites +
S. discolor +
S. rigoi
S. corsica +
S. bourgaeana +
S. cossoniana
S. gemmulosa +
S. cintrana
S. tenella (2) +
S. sedoides subsp. hohenwartii +
S. tenella (1) +
S. paradoxa +
S. berica +
S. presolanensis +
S. petraea +
S. arachnoidea +
S. androsacea +
S. seguieri +
S. aphylla +
S. depressa
S. irrigua +
S. adscendens +
S. italica +
S. osloensis (1)
S. biternata +
S. conifera (1)
S. conifera (2) +
S. haenseleri
0.07
SAXIFRAGA
Arachnoideae
Saxifraga
Androsaceae
Tridactylites


Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record
trnL-trnFPorphyrion  

Gymnopera S. spathularisS. umbrosa


Ligulatae  
TrachyphyllumtrnL-trnF

DISCUSSION
Major lineages of Saxifraga, morphological characteris-
tics and classification. —

Saxifraga 

Saxifraga

C
A
100/100
100/100
100/98
91/82
99/83
100/94
100/ 100
99/98
99/100
98/100
58/ -
91/-
95/97
77/ 77
74/-
74/-
81/62
72/67
61/ 61
94/87
65/-
95/ 78
62/-
65/-
92/-
52/-
76/-
68/-
65/-
81/57
78/-
51/62
72/70
96/97
86/83
68/53
94/ 81
84/82
68/53
66/-
65/63
63/-
76/-
85/90
82/-
91/60
88/87
70/65
94/ 85
100/100
100/100
100/100
98/64
78/66
76/53
79/71
79/50
62/-
94/ 82
86/89
100/100
99/99
100/98
100/99
100/100
75/95
88/96
100/100
100/100
100/100
100/100
100/100
90/96
100/100
100/100
99/86
55/ 63
86/80
86/81
100/94
53/73
70/86
96/87
PORPHYRION
LIGULATAE
TRACHYPHYLLUM
MESOGYNE
GYMNOPERA
COTYLEA
CYMBALARIA
Kabschia
Oppositifoliae
Florulentae
Squarrosae
Mutatae
S. stolitzkae +
S. andersonii +
S. matta-florida +
S. georgei +
S. alpigena +
S. lilacina +
S. pulvinaria +
S. afghanica +
S. duthiei +
S. roylei +
S. unguipetala +
S. scardica +
S. wendelboi +
S. marginata (1) +
S. alberti +
S. ferdinandi-coburgi +
S. marginata (2) +
S. kotschyi +
S. sancta +
S. marginata var. bubakii +
S. felineri +
S. diapensioides +
S. aretioides +
S. iranica +
S. charadzeae +
S. subverticillata +
S. vandellii +
S. tombeanensis +
S. caspica +
S. dinnikii +
S. caucasica +
S. colchica +
S. ruprechtiana +
S. pseudolaevis +
S. juniperifolia
S. scleropoda +
S. desoulavyi +
S. columnaris +
S. stribrnyi +
S. thessalica +
S. sempervivum +
S. federici-augusti subsp. grisebachii +
S. porophylla +
S. spruneri +
S. corymbosa +
S. luteoviridis +
S. oppositifolia (1)
S. oppositifolia subsp. asiatica (1) +
S. oppositifolia subsp. asiatica (2) +
S. oppositifolia subsp. oppositifolia +
S. oppositifolia subsp. smalliana
S. oppositifolia (2)
S. retusa subsp. augustana +
S. mutata +
S. aizoides +
S. squarrosa
S. caesia +
S. florulenta
S. valdensis (1)
S. cf. cotyledon +
S. valdensis (2) +
S. cotyledon +
S. cochlearis +
S. crustata +
S. hostii +
S. paniculata +
S. cartilaginea +
S. kolenatiana +
S. lingulata +
S. callosa +
S. bryoides +
S. aspera +
S. spathularis (1)
S. coriifolia +
S. taygetea +
S. rotundifolia +
S. spathularis (2) +
S. umbrosa +
S. hirsuta
S. hederacea (2)
S. hederacea (1)
S. cymbalaria (1) +
S. cymbalaria (2) +
S. sibthorpii +
S. cuneifolia +
S. blepharophylla
S. biflora +
S. rivularis +
S. hyperborea +
S. radiata (1) +
S. radiata (2) +
S. bracteata (1) +
S. bracteata (2) +
S. cernua (2) +
S. cernua (1)
S. carpatica +
S. sibirica +
S. granulifera +
S. flexuosa +
S. rivularis subsp. arctolitoralis +
0.07
S. cespitosa subsp. monticola
S. cespitosa subsp. cespitosa
S. terektensis
S. rosacea
S. hypnoides
S. continentales
S. cuneata
S. fragosoi
S. maderensis
S. portosancta
S. trifurcata
S. babiana
S. camposii var. leptophylla
S. canaliculata
S. latepetiolata (2)
S. latepetiolata (1)
S. dichotoma
S. carpetana
S. bulbifera
S. granulata
S. moschata
S. graeca
S. pentadactylis subsp. willkommiana
S. pentadactylis (1)
S. pentadactylis (2)
S. nevadensis
S. geranioides
S. magellanica
S. moncayensis
S. genesiana
S. vayredana
S. cebennensis
S. exarata
S. pontica
S. praetermissa
S. pubescens subsp. iratiana
S. intricata
S. hariotii
S. fragilis (2)
S. fragilis (1)
S. aquatica
S. pedemontana subsp. cymosa
S. reuteriana
S. globulifera (1)
S. globulifera (2)
S. erioblasta
S. trabutiana
S. wahlenbergii
S. osloensis (1)
S. tridactylites
S. discolor
S. rigoi
S. corsica
S. bourgaeana
S. cossoniana
S. gemmulosa
S. cintrana
S. tenella (2)
S. sedoides subsp. hohenwartii
S. tenella (1)
S. paradoxa
S. berica
S. presolanensis
S. petraea
S. arachnoidea
S. androsacea
S. seguieri
S. aphylla
S. depressa
S. irrigua
S. adscendens
S. italica
S. osloensis (2)
S. stolitzkae
S. andersonii
S. matta-florida
S. georgei
S. alpigena
S. lilacina
S. pulvinaria
S. afghanica
S. duthiei
S. roylei
S. unguipetala
S. scardica
S. wendelboi
S. marginata (1)
S. albertii
S. ferdinandi-coburgi
S. marginata (2)
S. kotschyi
S. sancta
S. marginata var. bubakii
S. felineri
S. diapensioides
S. aretioides
S. iranica
S. charadzeae
S. subverticillata
S. vandellii
S. tombeanensis
S. caspica
S. dinnikii
S. caucasica
S. colchica
S. ruprechtiana
S. pseudolaevis
S. juniperifolia
S. scleropoda
S. desoulavyi
S. columnaris
S. stribrnyi
S. thessalica
S. sempervivum
S. federici-augusti subsp. grisebachii
S. porophylla
S. spruneri
S. corymbosa
S. luteo-viridis
S. oppositifolia (1)
S. oppositifolia subsp. asiatica (1)
S. oppositifolia subsp. asiatica (2)
S. oppositifolia subsp. oppositifolia
S. oppositifolia subsp. smalliana
S. oppofitifolia (2)
S. retusa subsp. augustana
S. mutata
S. aizoides
S. squarrosa
S. caesia
S. florulenta
S. valdensis (1)
S. cf. cotyledon
S. valdensis (2)
S. cotyledon
S. cochlearis
S. crustata
S. hostii
S. paniculata
S. cartilaginea
S. kolenatiana
S. lingulata
S. callosa
S. bryoides
S. aspera
S. spathularis (1)
S. coriifolia
S. taygetea
S. rotundifolia
S. spathularis (2)
S. umbrosa
S. hirsuta
S. hederacea (2)
S. hederacea (1)
S. cymbalaria (1)
S. cymbalaria (2)
S. sibthorpii
S. saginoides
S. viscidula
S. aristulata
S. montana
S. pseudohirculus
S. peplidifolia
S. litangensis
S. yarlungzangboensis
S. przewalskii
S. hirculoides
S. tangutica
S. sinomontana
S. heleonastes
S. cf. hirculoides
S. tibetica
S. hirculus var. alpina
S. hirculus var. hirculus (1)
S. hirculus var. hirculus (2)
S. insolens
S. diversifolia var. haematophylla
S. egregia
S. hookeri
S. implicans
S. moorcroftiana
S. latifolia
S. lychnitis
S. isophylla
S. zimmermannii
S. parnassifolia
S. bergenioides
S. caveana
S. diversifolia
S. xiaozhongdianensis
S. kingdonii
S. kingiana
S. nanella
S. stella-aurea var. poyadena
S. gemmigera var. gemmuligera
S. contraria f. rubella
S. engleriana
S. unguiculata
S. aurantiaca
S. tatsienluensis
S. perpusilla
S. heterotricha
S. umbellatula var. umbellatula
S. signatella
S. umbellatula var. muricola
S. dingqingensis
S. umbellatula var. pectinata
S. serpyllifolia subsp. glutinosa
S. aleutica
S. serpyllifolia var. serpyllifolia
S. sanguinea
S. chrysantha
S. jaquemontiana
S. finitima
S. punctulata
S. sediformis
S. macrocalyx
S. platysepala
S. setigera
S. flagellaris subsp. crandallii
S. flagellaris subsp. crassiflagellata
S. flagellaris subsp. stenophylla
S. zhidoensis
S. mucronulata
S. mucronulatoides
S. pilifera
S. nigroglandulifera
S. nangxianensis
S. komarovii
S. consanguinea
S. angustata
S. hispidula (1)
S. hispidula (2)
S. strigosa var. ramosa
S. wallichiana
S. gouldii
S. brachypoda
S. gemmipara
S. filicaulis
S. substrigosa
S. eschscholtzii (1)
S. eschscholtzii (2)
S. firma (2)
S. funstonii (2)
S. funstonii (1)
S. nishidae
S. firma (1)
S. verspertina
S. taylori
S. sieversiana
S. korshinskii
S. lactea
S. bicuspidata
S. veitchiana
S. stolonifera
S. rufescens
S. mertensiana
S. nipponica
OUTGROUP
S. fortunei var. incisolobata
S. fortunei
S. cortusifolia
S. omolojensis (2)
S. anadyrensis (2)
S. anadyrensis (1)
S. spinulosa
S. omolojensis (1)
S. kruhsiana
S. stelleriana
S. cherlerioides
S. derbekii
S. bronchialis
S. bronchialis subsp. austromontana
S. caulescens
S. tricuspidata (1)
S. tricuspidata (2)
S. cherlerioides var. rebunshirensis
S. wardii
S. brunonis (1)
S. brunonis (2)
S. cuneifolia
S. blepharophylla
S. biflora
S. rivularis
S. hyperborea
S. radiata (1)
S. radiata (2)
S. bracteata (1)
S. bracteata (2)
S. cernua (2)
S. cernua (1)
S. carpatica
S. sibirica
S. granulifera
S. flexuosa
S. rivularis subsp. arctolitoralis
S. biternata
S. conifera (1)
S. conifera (2)
S. haenseleri
A
B
C


TAX ON
–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record
54/-
63/62
73/-
81/-
70/-
71/68
90/-
50/-
63/70
72/74
67/81
61/52
53/-
63/-
68/66
61/-
54/-
59/-
87/ 74
82/ 71
98/90
98/89
92/83
62/66
50/-
72/60
54/53
97/-
59/-
53/-
97/95
50/52
55/-
55/ -
53/56
84/-
78/84
100/96
100/99
64/74
59/62
84/88
84/85
100/97
99/95
100/95
100/95
100/100
100/100
100/100
100/86
93/91
99/100
100/100
100/100
100/100
57/56
100/-
100/100
99/100
99/100
100/99
100/100
100/99
65/95
83/74
68/100
59/86
50/96
81/99
83/ 92
91/92
93/91
94/97
100/100
100/90
100/100
S. saginoides +
S. viscidula +
S. aristulata +
S. sinomontana (1) +
S. pseudohirculus +
S. peplidifolia
S. litangensis +
S. yarlungzangboensis +
S. przewalskii +
S. hirculoides +
S. tangutica +
S. sinomontana (2) +
S. heleonastes +
S. cf. hirculoides +
S. tibetica +
S. hirculus var. alpina +
S. hirculus var. hirculus (1) +
S. hirculus var. hirculus (2) +
S. insolens
S. diversifolia var. haematophylla
S. egregia
S. hookeri
S. implicans +
S. moorcroftiana +
S. latifolia +
S. lychnitis +
S. isophylla +
S. zimmermannii +
S. parnassifolia +
S. bergenioides +
S. caveana +
S. diversifolia +
S. xiaozhongdianensis
S. kingdonii +
S. kingiana
S. nanella +
S. stella-aurea var. poyadena +
S. gemmigera var. gemmuligera +
S. contraria f. rubella +
S. engleriana +
S. unguiculata +
S. aurantiaca +
S. tatsienluensis
S. perpusilla +
S. heterotricha +
S. umbellulata var. umbellulata +
S. signatella +
S. umbellulata var. muricola +
S. dingqingensis
S. umbellulata var. pectinata
S. serpyllifolia subsp. glutinosa +
S. aleutica +
S. serpyllifolia subsp. serpyllifolia +
S. sanguinea
S. chrysantha +
S. jacquemontiana +
S. finitima +
S. punctulata
S. sediformis +
S. macrocalyx +
S. platysepala +
S. setigera
S. flagellaris subsp. crandallii +
S. flagellaris subsp. crassiflagellata +
S. flagellaris subsp. stenophylla +
S. zhidoensis
S. mucronulata +
S. mucronulatoides +
S. pilifera +
S. nigroglandulifera +
S. nangxianensis
S. komarovii +
S. consanguinea +
S. angustata +
S. hispidula (1)
S. hispidula (2) +
S. strigosa var. ramosa
S. wallichiana +
S. gouldii +
S. brachypoda +
S. gemmipara
S. filicaulis +
S. substrigosa +
S. eschscholtzii (1) +
S. eschscholtzii (2) +
S. firma (2) +
S. funstonii (2) +
S. funstonii (1) +
S. nishidae
S. firma (1) +
S. verspertina
S. taylorii +
S. sieversiana
S. korshinskyi
S. lactea +
S. bicuspidata
S. veitchiana +
S. nipponica +
OUTGROUP
S. stolonifera +
S. rufescens +
S. mertensiana
S. fortunei var. incisolobata +
S. fortunei +
S. cortusifolia +
S. omolojensis (2) +
S. anadyrensis (2) +
S. anadyrensis (1) +
S. spinulosa +
S. omolojensis (1) +
S. kruhsiana +
S. stelleriana
S. cherlerioides +
S. derbekii +
S. bronchialis +
S. bronchialis subsp. austromontana +
S. caulescens
S. tricuspidata (1) +
S. tricuspidata (2) +
S. cherlerioides var. rebunshirensis +
S. wardii +
S. brunonis (1)
S. brunonis (2) +
B
0.07
CILIATAE
SAXI-
FRAGELLA
PSEUDO-
CYMBALARIA
IRREGULARES
BRONCHIALES
S. cespitosa subsp. monticola
S. cespitosa subsp. cespitosa
S. terektensis
S. rosacea
S. hypnoides
S. continentales
S. cuneata
S. fragosoi
S. maderensis
S. portosancta
S. trifurcata
S. babiana
S. camposii var. leptophylla
S. canaliculata
S. latepetiolata (2)
S. latepetiolata (1)
S. dichotoma
S. carpetana
S. bulbifera
S. granulata
S. moschata
S. graeca
S. pentadactylis subsp. willkommiana
S. pentadactylis (1)
S. pentadactylis (2)
S. nevadensis
S. geranioides
S. magellanica
S. moncayensis
S. genesiana
S. vayredana
S. cebennensis
S. exarata
S. pontica
S. praetermissa
S. pubescens subsp. iratiana
S. intricata
S. hariotii
S. fragilis (2)
S. fragilis (1)
S. aquatica
S. pedemontana subsp. cymosa
S. reuteriana
S. globulifera (1)
S. globulifera (2)
S. erioblasta
S. trabutiana
S. wahlenbergii
S. osloensis (1)
S. tridactylites
S. discolor
S. rigoi
S. corsica
S. bourgaeana
S. cossoniana
S. gemmulosa
S. cintrana
S. tenella (2)
S. sedoides subsp. hohenwartii
S. tenella (1)
S. paradoxa
S. berica
S. presolanensis
S. petraea
S. arachnoidea
S. androsacea
S. seguieri
S. aphylla
S. depressa
S. irrigua
S. adscendens
S. italica
S. osloensis (2)
S. stolitzkae
S. andersonii
S. matta-florida
S. georgei
S. alpigena
S. lilacina
S. pulvinaria
S. afghanica
S. duthiei
S. roylei
S. unguipetala
S. scardica
S. wendelboi
S. marginata (1)
S. albertii
S. ferdinandi-coburgi
S. marginata (2)
S. kotschyi
S. sancta
S. marginata var. bubakii
S. felineri
S. diapensioides
S. aretioides
S. iranica
S. charadzeae
S. subverticillata
S. vandellii
S. tombeanensis
S. caspica
S. dinnikii
S. caucasica
S. colchica
S. ruprechtiana
S. pseudolaevis
S. juniperifolia
S. scleropoda
S. desoulavyi
S. columnaris
S. stribrnyi
S. thessalica
S. sempervivum
S. federici-augusti subsp. grisebachii
S. porophylla
S. spruneri
S. corymbosa
S. luteo-viridis
S. oppositifolia (1)
S. oppositifolia subsp. asiatica (1)
S. oppositifolia subsp. asiatica (2)
S. oppositifolia subsp. oppositifolia
S. oppositifolia subsp. smalliana
S. oppofitifolia (2)
S. retusa subsp. augustana
S. mutata
S. aizoides
S. squarrosa
S. caesia
S. florulenta
S. valdensis (1)
S. cf. cotyledon
S. valdensis (2)
S. cotyledon
S. cochlearis
S. crustata
S. hostii
S. paniculata
S. cartilaginea
S. kolenatiana
S. lingulata
S. callosa
S. bryoides
S. aspera
S. spathularis (1)
S. coriifolia
S. taygetea
S. rotundifolia
S. spathularis (2)
S. umbrosa
S. hirsuta
S. hederacea (2)
S. hederacea (1)
S. cymbalaria (1)
S. cymbalaria (2)
S. sibthorpii
S. saginoides
S. viscidula
S. aristulata
S. montana
S. pseudohirculus
S. peplidifolia
S. litangensis
S. yarlungzangboensis
S. przewalskii
S. hirculoides
S. tangutica
S. sinomontana
S. heleonastes
S. cf. hirculoides
S. tibetica
S. hirculus var. alpina
S. hirculus var. hirculus (1)
S. hirculus var. hirculus (2)
S. insolens
S. diversifolia var. haematophylla
S. egregia
S. hookeri
S. implicans
S. moorcroftiana
S. latifolia
S. lychnitis
S. isophylla
S. zimmermannii
S. parnassifolia
S. bergenioides
S. caveana
S. diversifolia
S. xiaozhongdianensis
S. kingdonii
S. kingiana
S. nanella
S. stella-aurea var. poyadena
S. gemmigera var. gemmuligera
S. contraria f. rubella
S. engleriana
S. unguiculata
S. aurantiaca
S. tatsienluensis
S. perpusilla
S. heterotricha
S. umbellatula var. umbellatula
S. signatella
S. umbellatula var. muricola
S. dingqingensis
S. umbellatula var. pectinata
S. serpyllifolia subsp. glutinosa
S. aleutica
S. serpyllifolia var. serpyllifolia
S. sanguinea
S. chrysantha
S. jaquemontiana
S. finitima
S. punctulata
S. sediformis
S. macrocalyx
S. platysepala
S. setigera
S. flagellaris subsp. crandallii
S. flagellaris subsp. crassiflagellata
S. flagellaris subsp. stenophylla
S. zhidoensis
S. mucronulata
S. mucronulatoides
S. pilifera
S. nigroglandulifera
S. nangxianensis
S. komarovii
S. consanguinea
S. angustata
S. hispidula (1)
S. hispidula (2)
S. strigosa var. ramosa
S. wallichiana
S. gouldii
S. brachypoda
S. gemmipara
S. filicaulis
S. substrigosa
S. eschscholtzii (1)
S. eschscholtzii (2)
S. firma (2)
S. funstonii (2)
S. funstonii (1)
S. nishidae
S. firma (1)
S. verspertina
S. taylori
S. sieversiana
S. korshinskii
S. lactea
S. bicuspidata
S. veitchiana
S. stolonifera
S. rufescens
S. mertensiana
S. nipponica
OUTGROUP
S. fortunei var. incisolobata
S. fortunei
S. cortusifolia
S. omolojensis (2)
S. anadyrensis (2)
S. anadyrensis (1)
S. spinulosa
S. omolojensis (1)
S. kruhsiana
S. stelleriana
S. cherlerioides
S. derbekii
S. bronchialis
S. bronchialis subsp. austromontana
S. caulescens
S. tricuspidata (1)
S. tricuspidata (2)
S. cherlerioides var. rebunshirensis
S. wardii
S. brunonis (1)
S. brunonis (2)
S. cuneifolia
S. blepharophylla
S. biflora
S. rivularis
S. hyperborea
S. radiata (1)
S. radiata (2)
S. bracteata (1)
S. bracteata (2)
S. cernua (2)
S. cernua (1)
S. carpatica
S. sibirica
S. granulifera
S. flexuosa
S. rivularis subsp. arctolitoralis
S. biternata
S. conifera (1)
S. conifera (2)
S. haenseleri
A
B
C


Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record
  
 matKrbcL trnL-trnF

      
Saxifraga



 






Section Irregulares. —


Diptera  Saxifraga
mertensiana 
  Boraphila  
Micranthes   
S. mertensiana   
Micranthes Heterisia
Saxifraga
Micranthes

Saxifraga 
 S. mertensiana
M. nelsoniana 
  S.  Micranthes   
S. Diptera
 IrregularesHeterisia
  Diptera
.
Irregulares 
Heterisia

Irregulares  


  
 
100/100
100/100
100/99 78/78
98/98
83/86
SAXIFRAGA
73/84
83/-
75/66
100/100
100/93
51/-
100/100
100/100
100/100
100/100
100/100
100/89
100/100 100/100
0.07
100/95
100/100
50/69 96/76
74/53
100/93
100/95
98/99
73/59
56/-
54/59
57/-
61/-
69/-
76/66
77/-
77/67
78/-
90/76
100/100
100/100
Chrysosplenium iowense
Micranthes brachypetala
Micranthes nudicaulis
Micranthes merkii
Micranthes tolmiei
Micranthes dicaricata
Micranthes tilingiana
Micranthes stellaris
Micranthes razshivinii
Micranthes micranthidifolia
Micranthes pensylvanica
Micranthes nivalis
Micranthes californica
Micranthes reflexa
Micranthes integrifolia
Micranthes calycina
Micranthes nelsoniana
Micranthes redofskyi
Micranthes ferruginea
Micranthes lyallii
Micranthes melanocentra
Peltoboykinia tellimoides
Astilboides tabularis
Mukdenia rossii
Oresitrophe rupifraga
Mitella nuda
Darmera peltata
Rodgersia pinnata
Tiarella polyphylla
Conimitella williamsii
Elmera racemosa
Bensoniella oregona
Heuchera micrantha
Tellima grandiflora
Tolmiea menziesii
Lithophragma trifoliatum
Tanakaea radicans
Leparrhena pyrolifolia
Suksdorfia violacea
Sullivantia oregana
Telesonix heucheriformis
Jepsonia parryi
Saxifragopsis fragarioides
Astilbe microphylla
Saniculiphyllum guangxiense
Itea virginica
Pterostemon rotundifolius
Cascadia nuttallii (2)
Cascadia nuttallii (1)
Saxifragodes albowiana
S. cespitosa subsp. monticola
S. cespitosa subsp. cespitosa
S. terektensis
S. rosacea
S. hypnoides
S. continentales
S. cuneata
S. fragosoi
S. maderensis
S. portosancta
S. trifurcata
S. babiana
S. camposii var. leptophylla
S. canaliculata
S. latepetiolata (2)
S. latepetiolata (1)
S. dichotoma
S. carpetana
S. bulbifera
S. granulata
S. moschata
S. graeca
S. pentadactylis subsp. willkommiana
S. pentadactylis (1)
S. pentadactylis (2)
S. nevadensis
S. geranioides
S. magellanica
S. moncayensis
S. genesiana
S. vayredana
S. cebennensis
S. exarata
S. pontica
S. praetermissa
S. pubescens subsp. iratiana
S. intricata
S. hariotii
S. fragilis (2)
S. fragilis (1)
S. aquatica
S. pedemontana subsp. cymosa
S. reuteriana
S. globulifera (1)
S. globulifera (2)
S. erioblasta
S. trabutiana
S. wahlenbergii
S. osloensis (1)
S. tridactylites
S. discolor
S. rigoi
S. corsica
S. bourgaeana
S. cossoniana
S. gemmulosa
S. cintrana
S. tenella (2)
S. sedoides subsp. hohenwartii
S. tenella (1)
S. paradoxa
S. berica
S. presolanensis
S. petraea
S. arachnoidea
S. androsacea
S. seguieri
S. aphylla
S. depressa
S. irrigua
S. adscendens
S. italica
S. osloensis (2)
S. stolitzkae
S. andersonii
S. matta-florida
S. georgei
S. alpigena
S. lilacina
S. pulvinaria
S. afghanica
S. duthiei
S. roylei
S. unguipetala
S. scardica
S. wendelboi
S. marginata (1)
S. albertii
S. ferdinandi-coburgi
S. marginata (2)
S. kotschyi
S. sancta
S. marginata var. bubakii
S. felineri
S. diapensioides
S. aretioides
S. iranica
S. charadzeae
S. subverticillata
S. vandellii
S. tombeanensis
S. caspica
S. dinnikii
S. caucasica
S. colchica
S. ruprechtiana
S. pseudolaevis
S. juniperifolia
S. scleropoda
S. desoulavyi
S. columnaris
S. stribrnyi
S. thessalica
S. sempervivum
S. federici-augusti subsp. grisebachii
S. porophylla
S. spruneri
S. corymbosa
S. luteo-viridis
S. oppositifolia (1)
S. oppositifolia subsp. asiatica (1)
S. oppositifolia subsp. asiatica (2)
S. oppositifolia subsp. oppositifolia
S. oppositifolia subsp. smalliana
S. oppofitifolia (2)
S. retusa subsp. augustana
S. mutata
S. aizoides
S. squarrosa
S. caesia
S. florulenta
S. valdensis (1)
S. cf. cotyledon
S. valdensis (2)
S. cotyledon
S. cochlearis
S. crustata
S. hostii
S. paniculata
S. cartilaginea
S. kolenatiana
S. lingulata
S. callosa
S. bryoides
S. aspera
S. spathularis (1)
S. coriifolia
S. taygetea
S. rotundifolia
S. spathularis (2)
S. umbrosa
S. hirsuta
S. hederacea (2)
S. hederacea (1)
S. cymbalaria (1)
S. cymbalaria (2)
S. sibthorpii
S. saginoides
S. viscidula
S. aristulata
S. montana
S. pseudohirculus
S. peplidifolia
S. litangensis
S. yarlungzangboensis
S. przewalskii
S. hirculoides
S. tangutica
S. sinomontana
S. heleonastes
S. cf. hirculoides
S. tibetica
S. hirculus var. alpina
S. hirculus var. hirculus (1)
S. hirculus var. hirculus (2)
S. insolens
S. diversifolia var. haematophylla
S. egregia
S. hookeri
S. implicans
S. moorcroftiana
S. latifolia
S. lychnitis
S. isophylla
S. zimmermannii
S. parnassifolia
S. bergenioides
S. caveana
S. diversifolia
S. xiaozhongdianensis
S. kingdonii
S. kingiana
S. nanella
S. stella-aurea var. poyadena
S. gemmigera var. gemmuligera
S. contraria f. rubella
S. engleriana
S. unguiculata
S. aurantiaca
S. tatsienluensis
S. perpusilla
S. heterotricha
S. umbellatula var. umbellatula
S. signatella
S. umbellatula var. muricola
S. dingqingensis
S. umbellatula var. pectinata
S. serpyllifolia subsp. glutinosa
S. aleutica
S. serpyllifolia var. serpyllifolia
S. sanguinea
S. chrysantha
S. jaquemontiana
S. finitima
S. punctulata
S. sediformis
S. macrocalyx
S. platysepala
S. setigera
S. flagellaris subsp. crandallii
S. flagellaris subsp. crassiflagellata
S. flagellaris subsp. stenophylla
S. zhidoensis
S. mucronulata
S. mucronulatoides
S. pilifera
S. nigroglandulifera
S. nangxianensis
S. komarovii
S. consanguinea
S. angustata
S. hispidula (1)
S. hispidula (2)
S. strigosa var. ramosa
S. wallichiana
S. gouldii
S. brachypoda
S. gemmipara
S. filicaulis
S. substrigosa
S. eschscholtzii (1)
S. eschscholtzii (2)
S. firma (2)
S. funstonii (2)
S. funstonii (1)
S. nishidae
S. firma (1)
S. verspertina
S. taylori
S. sieversiana
S. korshinskii
S. lactea
S. bicuspidata
S. veitchiana
S. stolonifera
S. rufescens
S. mertensiana
S. nipponica
OUTGROUP
S. fortunei var. incisolobata
S. fortunei
S. cortusifolia
S. omolojensis (2)
S. anadyrensis (2)
S. anadyrensis (1)
S. spinulosa
S. omolojensis (1)
S. kruhsiana
S. stelleriana
S. cherlerioides
S. derbekii
S. bronchialis
S. bronchialis subsp. austromontana
S. caulescens
S. tricuspidata (1)
S. tricuspidata (2)
S. cherlerioides var. rebunshirensis
S. wardii
S. brunonis (1)
S. brunonis (2)
S. cuneifolia
S. blepharophylla
S. biflora
S. rivularis
S. hyperborea
S. radiata (1)
S. radiata (2)
S. bracteata (1)
S. bracteata (2)
S. cernua (2)
S. cernua (1)
S. carpatica
S. sibirica
S. granulifera
S. flexuosa
S. rivularis subsp. arctolitoralis
S. biternata
S. conifera (1)
S. conifera (2)
S. haenseleri
A
B
C


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–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record
Fig. 5.
SaxifragaSaxifraga trnL-
trnF
   
trnL-trnF
S. rosacea
S. cespitosa subsp. cespitosa
S. terektensis
S. cespitosa subsp. monticola
S. hypnoides
S. continentalis
S. cuneata
S. fragosoi
S. maderensis*
S. portosanctana*
S. trifurcata
S. babiana
S. camposii var. leptophylla
S. canaliculata
S. latepetiolata (2)*
S. latepetiolata (1)
S. dichotoma
S. carpetana*
S. bulbifera
S. granulata
S. moschata
S. graeca
S. pentadactylis subsp. willkommiana
S. pentadactylis (1)
S. pentadactylis (2)
S. nevadensis
S. geranioides
S. magellanica
S. moncayensis
S. genesiana*
S. vayredana
S. cebennensis
S. exarata
S. pontica*
S. praetermissa
S. pubescens subsp. iratiana
S. intricata*
S. hariotii*
S. fragilis (2)*
S. fragilis (1)
S. aquatica*
S. pedemontana subsp. cymosa
S. reuteriana*
S. globulifera (1)
S. globulifera (2)
S. erioblasta
S. trabutiana*
S. wahlenbergii
S. osloensis (2)
S. tridactylites
S. discolor*
S. rigoi*
S. corsica
S. bourgaeana
S. cossoniana*
S. gemmulosa
S. cintrana*
S. tenella (2)
S. sedoides subsp. hohenwartii
S. tenella (1)
S. paradoxa
S. berica
S. presolanensis
S. petraea
S. arachnoidea
S. androsacea
S. seguieri
S. aphylla
S. depressa*
S. irrigua
S. adscendens
S. italica
S. osloensis (1)
S. biternata
S. conifera (1)
S. conifera (2)*
S. haenseleri*
S. cespitosa subsp. cespitosa
S. magellanica
S. cespitosa subsp. monticola
S. moschata
S. continentalis
S. canaliculata
S. hypnoides
S. fragosoi
S. trifurcata
S. babiana
S. camposii var. leptophylla
S. cuneata
S. latepetiolata (1)
S. dichotoma
S. bulbifera
S. granulata
S. terektensis
S. graeca
S. pentadactylis (1)
S. pentadactylis (2)
S. nevadensis
S. geranioides
S. rosacea
S. moncayensis
S. vayredana
S. cebennensis
S. exarata
S. praetermissa
S. pubescens subsp. iratiana
S. fragilis (1)
S. pedemontana subsp. cymosa
S. globulifera (1)
S. globulifera (2)
S. erioblasta
S. wahlenbergii
S. adscendens
S. osloensis (1)
S. corsica
S. bourgaeana
S. gemmulosa
S. tenella (2)
S. aphylla
S. tenella (1)
S. paradoxa
S. berica
S. sedoides subsp. hohenwartii
S. petraea
S. arachnoidea
S. androsacea
S. seguieri
S. presolanensis
S. irrigua
S. osloensis (2)
S. italica
S. tridactylites
S. biternata
S. conifera (1)
99
-
95
88
100
99
95
83
99
99
89
88
88
82
100
100
100
100
100
91
100
98
100
99 100
100
100
100
100
100
100
100
100
100
99
99
100
100
100
100
98
100
99
98
98
97
100
100
100
100
100
100
100
100
100
99
100
100
100
100
90
88
91
97
100
-
98
-
92
82
92
87
97
93
97
93
99
98
87
75
88
76
96
-
88
84
85
84
88
90
100
64
100
100
91
89
89
90
97
95
99
96
100
64
Nuclear ribosomal ITS
DNA sequence data
Plastid trnL-trnF
DNA sequence data
ML bootstrap values
MP bootstrap values


Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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–
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

Saxifraga mertensiana
Irregulares
 .  
   
Saxifraga
 
Mesogyne 
 Micranthes

 Saxifraga mertensiana
 Irregulares
   
S. fortuneiS. stoloniferaS. rufescens
S. nipponica
Saxifraga 

Micranthes Saxifragodes
Astilboides Darmera Rodgersia 
  Peltoboykinia 



. Irregulares


  Porphyrion
Ligulatae
Section Saxifragella —
Saxifraga bicuspidata



 Saxifragella

Saxifraga
 

Saxifraga  .
 
Saxifragella



 


Section Pseudocymbalaria. —
Saxifraga sieversiana
 S. korshinskyi 
  
Micranthes

  
 
Saxifraga


S. lactea

 


S. sieversianaS. korshinskyi

Saxifraga



Saxifraga

Micranthes



korshinskyiS. lactea

sieversiana



Saxifraga sieversiana
 S. selemdzhensis 
   Pseudocymba-
lariaS. korshinskyi   Komarovia
Saxifraga lactea
Lacteae
Tridactylites 
S. lactea
  Tridactylites 


Tridactylites

    
S. lacteaS. sieversianaS. korshinsk yi
   


S. sieversiana

S. korshinskyiS. lactea

Section Bronchiales. —














  

  Trachyphyllum
   



S. aspera S. bry-
oides

Trachyphyllum  
Bron-
chiales


Bronchiales
Trachyphyllum

  Saxifraga. 


  



Bronchiales
  
       
    
 

Porphyrion

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–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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
 Bronchiales  


Section Ciliatae. —
 Saxifraga
 
   trnL-trnF 
   

   

S. hirculus S. flagellaris 
  

S. chrysantha
 S. serpyllifolia  
S. platysepala
S. eschscholtzii  
S. aleutica

  
 

Ciliatae 
  

     
   
    


        Cilia-
tae   


Ciliatae
 
S. lychnitis
S. nigroglandulifera

 Ciliatae       
             
     
S. eschscholtzii

S. filicaulis S. gemmip-
araS. hispidula
Gemmiparae   

S. angustata S. flagellarisS. seti-
gera
 
  
Flagellares Saxifraga
eschscholtziiS. zhidoensis
Hemisphaericae  


 Ciliatae



Saxifraga aleuticaS. chrysanthaS. ser-
pyllifolia

 

Serpyllifoliae  
Rosulares
Sediformes
    
 





 S. sanguineaRosulares
Serpyllifoliae
trnL-trnFS. sanguinea
   

    
   

S. aristulata
S. diversifolia 

Hirculoideae



Saxifraga hirculus
   

Section Cymbalaria. —
 



S. cymbalariaS. hederacea S. sibthorpii
S. hederifolia
 


S. hederacea 
Saxifraga
SaxifragaCymbalaria
   


 Cymbalaria  
   



Section Cotylea. —
       
 Cot ylea S. taygetea


Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record
 
S. rotundifolia 

S. coriifolia
S. rotundi-
folia
  
 Cotylea


 

Section Gymnopera. —


  S. cuneifoliaS. hir-
sutaS. spathularis S. umbrosa


  Cotylea Ligulatae
Mesogyne PorphyrionTrachyphyllum 
trnL-trnF 
matK-trnK

     Gymnopera 
  



    
Cotylea 


Gymnopera

Section Mesogyne. —

Mesog yne




  
Mesogyne


    
S. cernua

S. mertensiana Irregulares
  



Mesog yne 
  S. carpat-
icaS. cernuaS. granulifera S. sibir-
icaS. rivularis
S. bracteata  S. f lexuosa  S. hyperborea
 S. rivularis  
S. radiata

Section Trachyphyllum. —
 Bron-
chiales   S. asperaS. bryoides
   Trachyphyllum

 Fibrosophyllae
Saxifraga asperaS. bryoides
   


   Trachyphyllum 
Ligulatae
Porphyrion
S. aspera
S. bryoides
 LigulataePorphyrion

S. cotyledon S. hostii    S. paniculata

Section Ligulatae. —

Ligulatae
Saxifraga paniculata
 

S. florulenta S. mutata
  
matKPorphyrion
.S. florulentaS. mutata
Ligulatae
 
  

Ligulatae


   S. cotyledon 

S. florulentaS. mutata


Section Porphyrion. —
 

S. oppositifolia 

Porphyrion   

 
S. aizoidesXanthizoon
  Ligulatae
S. florulentaS. mutata 
Florulentae  Mutatae

Porphyrion 

  

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–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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       
OppositifoliaeLigulataePor-
phyrion   

Porphyrion
 


S. caesiaS. squar-
rosa   S. aizoides S. mutata    
     S. caesia
S. aizoides S. squarrosa  S. aizoides 
S. caesiaS. mutata
  
    
Saxifraga caesia 
S. squarrosa



 
   
Squarrosae

 Saxifraga aizoidesS. mutata
Porphyrion
XanthizoonLigula-
tae
matK


 
Saxifraga aizoidesS. mutata
Mutatae



 

Mutatae

S. aizoidesS. mutata


S . biflora
S. blepharophylla S. oppositi-
folia  S. retusa
  Oppositifoliae 
 
S. oppositifolia 

 
   

S. oppositi-
folia  



 Saxifraga florulenta

Florulen-
tae


    
Porphyrion     
S. florulenta  

S. florulenta  
Porphyrion
Ligulatae
Porphyrion 

       
  
S. sancta
S. juniperifoliaB


  


Kabschia    
Kabschia

Squarrosae

Kabschia


Saxi-
fraga spruneri
Kabschia 
  


S. stribrnyi
S. sempervivum 

 
  
  Mediae   

S. caucasicaS. pseudolaevis

    



Juniperifoliae 


  


Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record


S. albertiS. roylei




Section Saxifraga. —
  
S. granulata
Saxifraga 


   


Saxifraga 

 

Saxifraga
SaxifragaSaxifraga

  


 Saxifraga 
S. adscendensS. discolorS. oslo-
ensis  S. tridactylites 
Tridactylites


S. osloensis
S. adscendensS. tridactylites
    
 

S. wahlenbergii   
trnL-trnF 

  Tridactylites
   

Tridactylites

Androsaceae

S. androsaceaS. depressa  S. italica
    S. seguieri   



S. androsacea

S. irrigua


Saxifraga
irrigua
 Androsaceae

S. aquaticaS. latepetiolata
Saxifraga


Arachnoideae    


  trnL-trnF







S. facchinii
  

Saxifraga

Saxifraga

    


  
Saxifraga
SaxifragaS. conifera 
 

S. conifera   
 
Saxifraga



S. bourgae
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ana S. gemmulosa

S. bulbiferaS. carpetanaS. dichotoma
S. graeca 
S. globuliferaS. reuteriana
S. trabutiana
S. maderensisS. portosanctana
  

 S. babiana
S. trifurcata
  
   
S. exarataS. moschata
S. cespitosacespitosamon-
ticola  


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TAX ON
–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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
 



Saxi-
fraga
   

Unplaced taxa. —
Saxifraga odontophylla
 Boraphila 

Micranthes 
   
Micranthes
S. Odontophylla.
Micranthes
SSaxifraga
  
S. odontophylla
S. sieversiana S. korshinskyi
S. odontophylla S. sibirica
Mesog yne. Saxifraga
odontophylla

S. decus-
sata S. monantha S. nana
 Saxi-
fraga Tetrameridium
 
Tetrameridium
Oppositifoliae 
Tet r a m e r idium

Delineation of Saxifraga. —
Cascadia nuttallii
Saxifraga nuttallii
S. tridac-
tylitesS. petraeaTridactylites
Cascadia
Saxifraga

 Cascadia
Saxifraga
Saxifragodes

trnL-trnFSaxifragodes albowiana

Hybridisation, hybrid speciation and allopolyploidy in
Saxifraga. —
 
  
Saxifraga


   

Saxifraga 


       

Saxifraga
S. moschata 
S. osloensisS. adscendens × S. tridactylites
S. opdalensisS. svalbardensis
 S. cernua × S. rivularis

Saxifraga






 



S. osloensis

trnL-trnF 
Saxifraga osloensis
 trnL-trnF S. adscen-
densS. tridactylites
  
S. adscendens 
 S. tridactylites  
S. adscendens
S. tridactylites  
        
Saxifraga
     
trnL-trnF
 
   
  
    

 
    



 
     Saxifraga 


   
    
  Bronchiales 
Gymnopera Ligulatae 
 Cymbalaria 

 



Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record
Abbott, R .J., Smith, L.C., Milne, R.I., Crawford, R.M.M., Wolff,
K. & Balfour, J. 
Science

Akiyama, S. & Gornall, R.J. Saxifraga

 Flora of Nepal: Saxifragaceae

 

Baker, W. J., Norup, M.V., Clarkson, J.J., Couvreur, T.L.P., Dowe,
J.L., Lewis, C.E., Pintaud, J.- C., Savolainen, V., Wilmot, T.
& Chase, M.W.   
Ann. Bot. (Oxford)


Ball, T. Saxifraga. Bot. Zei-
tung (Berlin) 
Beatty, G.E., Barker, L., Chen, P.-P., Kelleher, C.T. & Provan, J.
   Saxifraga
hirsuta Saxifraga
spathularis    Ann.
Bot. (Oxford)
 
Böcher, T.W. Saxifraga nathorstii
 S. aizoidesS. oppositifolia. Meddel. Grøn-
land Biosci.
Brewbaker, J.L. . 
. Amer.
J. Bot. : –

Brochmann, C. & Håpnes, A. 
Saxifraga
  
SaxifragaS. wahlenbergii

S. adscendens S. discolor S. osloensisS. tri-
dactylites Tridactylites . 
trnL-trnF S. wahlenbergii

S. androsaceaS. seguieri
Androsaceae 
     S. wahlenbergii 
S. praetermissa
S. adscen-
dens S. androsacea S. exarataS. lactea S. styri-
aca
Perdurantes
S. wahlenbergii

n =
S. wahlenbergiiS. tridac-
tylitesnS. adscendensnS. praetermissa
n
xSaxifraga androsacea
n =

          
Saxifraga     S. aphylla S. presolanensis
S. sedoides hohenwartii                
Arachnoideae

S. cespitosa S. continenta-
lis   S. graeca S. granulata S. hyp-
noides S. moschata S. rosaceaS. terektensis
Saxifraga

S. moschataS. rosa-
cea   





NEW COMBINATIONS

 Saxifraga


SaxifragaSaxifragaAndrosaceae
    stat. nov.
Saxifraga  Dactyloides Holo-
phyllaeAndrosaceae
 
   
Saxifraga androsacea L.
SaxifragaSaxifragaArachnoideae
stat. nov. Saxi-
fragaNephrophyllum
Arachnoideae
  
Saxifraga arachnoidea
SaxifragaPorphyrion Squarrosae
stat. nov.
SaxifragaKabschia Squarrosae


Saxifraga squarrosa 
ACKNOWLEDGEMENTS
       
     

   
    

           
     


LITERATURE CITED


TAX ON
–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record
S. svalbardensis Bot. J. Linn.
Soc. 
Brochmann, C., Nilsson, T. & Gabrielsen, T.M.   

  Saxifraga osloensis
Acta Univ. Upsal., Symb. Bot. Upsal.
Brochmann, C., Xiang, Q.-Y., Brunsfeld, S.J., Soltis, D.E. & Soltis,
P. S . Saxifraga
S. svalbardensis 
Amer. J. Bot. 

Brouillet, L. & Elvander, P.E.    Saxifraga 
      Flora of North
America north of Mexico  Magnoliophyta: Paeoniaceae to
Ericaceae. 
Calder, J.A. & Savile, D.B.O. Saxi-
fragaTrachyphyllum   Brittonia 

Conti, E., Soltis, D.E., Hardig, T.M. & Schneider, J. 
SaxifragaLigula-
tae 
Molec. Phylogen. Evol. 

DeChaine, E.G. 
Saxifraga

Bronchiales
Rhodora
DeChaine, E.G., Anderson, S.A., McNew, J.M. & Wendling, B.M.

Saxifraga

Trachyphyllum

PLoS
ONE

Deng, J., Drew, B.T., Mavrodiev, E.V., Gitzendanner, M.A., Soltis,
P.S. & Soltis, E.D. 
   
Molec. Phylogen. Evol.

 
Eidesen, P.B., Müller, E., Lettner, C., Alsos, I.G., Bender, M.,
Kristiansen, M., Peeters, B., Postma, F. & Verweij, K.F. 
  

Saxifraga oppositi-
folia

Polar Res.


Elven, R., Murray, D.F., Razzhivin, V.Y. & Yurtsev, B.A. (eds.) 

 


Engler, H.G.A.  .
Die natürlichen Pflanzenfamilien
Podostemaceae–Platanaceae
Engler, H.G.A. & Irmscher, E.  – Saxi-
fragaDas Pflanzenreich 

Engler, H.G.A. & Irmscher, E.  – Saxifraga
Das Pflanzenreich 

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Ciliatae psbA-trnHtrnL-F 
Ta xon 
 
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L. Bot. J. Linn. Soc.

Gornall, R.J. Saxifraga
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
Halda, J.J.     Porphyrion
Acta Mus. Richnov. Sect. Nat.
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 Saxifraga oppositifolia 
Amer. J. Bot. 


Horný, R., Webr, K.M. & Byam-Grounds, J. Porophyllum saxi-
frages
Johnson, A.M. Saxifraga nutallii. Amer. J. Bot

Jørgensen, M.H. Evolutionary lineages and species delimitation
in the  complex.
         


Jørgensen, M.H., Elven, R., Tribsch, A., Gabrielsen, T.M., Stedje,
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Saxifraga rivularis Syst. Bot.

Kaplan, K. 
   Saxifraga
Biblioth. Bot.
Kim, Y.-D., Kim, S.-Y., Shin, H. &
Kim, Y. D.

Astilbe
Korean J. Pl. Taxon
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SaxifragaWiss. Z. Pädagog.
Hochschule Potsdam, Math.-Naturwiss. Reihe

Köckinger, H. Saxifraga styriaca    

Phyton (Horn)


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Lithophragma 
 Syst. Bot.
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
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LagotisTaxo n 

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  
Science

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SaxifragaTriplinerviumJ. Syst. Evol.
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McGregor, M. S a xi f ra ges: A de f initive gui de to the 2000 sp eci es
hybrids and cultivars.


Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record
McGregor, M. & Harding, W. Saxifrages: The complete list of
species.
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    
Proceedings of the Gateway Computing Environments
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    Rev. Cytol.
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
MitellaMolec. Biol. Evol.
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Mitella
Molec. Phylogen. Evol.
  
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blicae Popularis SinicaeSaxifragaceae (1). 

Pan, J., Gornall, R. & Ohba, H. Saxifraga
Flora of China Brassic-
aceae–Saxifragaceae. 

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SaxifragisActa Soc. Bot. Poloniae

Pawłowska, S.      Saxifraga
wahlenbergiiS. perdurans  Fragm. Florist. Geobot.

Pfanzelt, S., García, C. & Mar ticorena, A.  
  Check
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Leitch, A.R., Leitch, I.J., Bennett, M.D., Soltis, P.S.
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Soltis,
D.E.
 
Tragopogon 

Amer. J. Bot.

 
Pirie, M.D.
    
 Ta xon 

Prieto, J.A.F., Arjona, J.M., Sanna, M., Pérez , R. & Cires, E. 
Micranthes
J. Pl. Res.
  
Rauscher, J.T., Doyle, J.J. & Brown, A.H.D. 
 
 Glycine tomentella  
Genetics
Schneider, J., Döring, E., Hilu, K.W. & Röser, M. 

matK trnKTa x o n

Senters, A.E. & Solt is, D.E. Ribes
  Taxo n 

Soltis, D.E.            The
families and genera of vascular plants
 
Soltis, D.E. & Soltis, P.S. 

Biological approaches and evolu-
tionar y trends in plants
  
Soltis, D.E., Johnson, L.A. & Looney, C. 
 Boykinia
Syst. Bot. 
Soltis, D.E., Kuzoff, R.K., Mort, M.E., Zanis, M., Fishbein, M.,
Hufford, L., Koontz, J. & Arroyo, M.K.   

Ann. Missouri Bot.
Gard. 
Soltis, D.E., Tago-Nakazawa, M., Xiang, Q.Y., Kawano, S., Murata,
J., Wakabayashi, M. & Hibsch-Jetter, C. 
Chrysosplenium
matKAmer. J. Bot

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B.C., Solt is, P.S., Burleigh, J.G. & Rubio De Casas, R. 
        
Amer. J. Bot.

Soltis, P.S. & Soltis, D.E.     
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
 
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Bioinformatics 

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 Saxifraga opdalensis  S. sval-
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
Swofford, D.L.   

Taberlet, P., Gielly, L., Pautou, G. & Bouvet, J.    
  

Pl. Molec. Biol.


Tkach, N., Röser, M. & Hoffmann, M.H. 
Micranthes
Bot. J. Linn. Soc. 

Vargas, P. Saxifraga

Flora iberica Ebenaceae-Saxifragaceae.

Vargas, P.  Saxifraga  Saxifraga
     Pl. Syst. Evol.

Vargas, P.  Saxi-
fraga pentadactylis
Nordic J. Bot. 
 
Vargas, P.  
Taxon  

Vargas, P. & Nieto Feliner, G. Saxi-
fraga Ceratophyllae Pl. Syst. Evol. 

Vargas, P., Morton, C.M. & Jury, S.L. 
 Saxifraga

Amer. J. Bot.

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TAX ON
–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record
Wakabayashi, M. & Ohba, H.     
Saxifraga. Bull. Univ. Mus. Univ. Tokyo


Webb, D.A.  Saxifraga 
 
Flora
europaea Psilotaceae to Platanaceae

Webb, D.A. & Gornall, R. J. Saxifrages of Europe: With notes
on African American and some Asiatic species
Westergaard, K.B., Jørgensen, M.H., Gabrielsen, T.M., Alsos, I.G.
& Brochmann, C. 
Saxifraga rivularis
J. Biogeogr.
  
White, T.J., Bruns, T., Lee, S. & Taylor, J. 


PCR protocols: A guide to methods and applications.

  
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Winkler, M., Tribsch, A., Schneeweiss, G.M., Brodbeck, S., Gugerli,
F., Holderegger, R., Abbott, R. J. & Schönswetter, P. 

Saxifraga oppositifoliaMolec. Ecol.
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    
Saxifraga
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Byull. Moskovsk . Obshch. Isp. Prir. Otd. Biol. 
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MicranthesDip-
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Zhmylev, P.Y. Saxifraga 
  Porphyrion 
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Zhmylev, P.Y.    
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SaxifragaByull. Moskovsk. Obshch.
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Zhmylev, P.Y. Rod L. (Saxifragaceae): Biomorfologia,
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Zhmylev, P.Y. & Khokhryakov, A. Tra -
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SaxifragaTrachyphyllum  Novosti Sist.
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Zhou, J., Gong, X., Downie, S.R. & Peng, H. 


rpl16r ps16 Molec. Phylogen. Evol
  
Zhu, W.-D., Nie, Z.-L., Wen, J. & Sun, H. 
Astilbe
Bot. J. Linn. Soc.
 
Appendix 1.


trnL-trnF

OUTGROUP ( Iteaceae): Itea virginica Pterostemon rotundifolius 
SAXIFRAGACEAE: Astilbe microphylla 
Astilboides tabularis Bensoniella oregona 
Cascadia nuttallii 
Chrysosplenium iowense Coni-
mitella williamsii Darmera peltata 
Elmera racemosa Heuchera micrantha 
Jepsonia parryi 
Leptarrhena pyrolifolia Lithophragma


Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
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–
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trifoliatum Micranthes brachypetala 
M. californica M. calycina  
M. divaricata   M. ferruginea    
M. integrifolia M. lyallii 
LM654452. M. melanocentra M. merkii 
M. micranthidifolia  M. nelsoniana  
M. nivalis  M. nudicaulis   
M. pensylvanica M. razshivinii 
M. redofskyi M. reflexa 
 M. stellaris  M. tilingiana 
M. tolmiei Mitella nuda
 Mukdenia rossii   
Oresitrophe rupifraga Peltoboykinia tellimoides 
Rodgersia pinnata Saniculiphyllum guangxiense
Saxifraga adscendens M. Röser 9710  LN812341LN812575.
S. afghanica Wu & al. 5030 LN812342LN812576. S. aizoides  
N. Tk ach  LN812343LN812577. S. alberti G. Lazkov & N.V. Kente-
baeva  LN812344LN812578. S. aleutica S. Talbot & S.L. Talbot 489 LN812345LN812579. S. alpigena  
N. Tkach 225 & M. Röser LN812346LN8125 80. S. anadyrensis 
Yu.P. Kozhevni ko v LN812347LN812581A.A. Korobkov & N.A. Sek retare va LN812348LN812582. S. andersonii
N. Tkach 228 & M. Röser  LN812349LN812583. S. androsacea M. Röser
6935    LN812350LN812584. S. angustata G. & S. Miehe & U. Wündisch 94-404-14 LN812351LN812585.
S. a phylla M. Röser 7045 LN812352LN812586. S. aquatica S. arachnoidea
 M. Röser 10259   LN812353LN812587. S. aretioides    
N. Tkach 209 & M. Röser    LN812354LN81258 8. S. aristulata  G. & S . Miehe 99-134-20  LN812355
LN812589. S. aspera M. Röser 5839   LN812356LN812590. S. aurantiaca G. & S. Miehe & U. Wün-
disch 94-339-12c LN812357LN812591. S. babiana   N. Tkach 210 &
M. Röser  LN812358LN812592. S. bergenioides G. & S. Miehe 00-357-03 LN812359LN812593. S. berica 
 H. Meltesics LN812360LN812594. S. bicuspidata 
S. biflora M. Röser 1430 LN812361LN812595. S. biternata 
 N. T kach 260 & M. Röser   L N812362LN812596. S. blepharophylla   
 S. bourgaeana M. Röser 4682 LN812364LN812598. S. brachypoda G. & S. Miehe
00 -385-10 LN812365LN812599. S. bracteata S. Kharkevich & V. Barkalov 423 LN812366LN812600
T. Kel so J. Flock & M. Colson 235 LN812367T. Derviz-Sokolovskaya 251LN812601. S. bronchialis  bron-
chialisT.G. Leonova LN812369LN812603. S. bronchialis  austromontana 
N. Tkach 259 & M. Röser   LN812368LN812602. S. brunonis  
G. & S. Miehe 00-293-01 LN812370LN812604. S. bryoides M. Röser 9602
   LN812371 LN812605. S. bulbifera M. Röser 9124    LN812372 LN812606. S. caesia 
M. Röser 10264     LN812373 LN812607. S. callosa  N. Tkach 67   
LN8 12374LN812608. S. camposii   leptophylla A. Seg ura Zubizarreta 
LN812375LN812609. S. canaliculata N. Tkach 211 & M. Röser  
LN812 376LN812610. S. carpatica M. Röser 9676 LN812377LN812611. S. carpetana  
 S. cartilaginea    A. Patudin  LN812378 LN812612. S. caspica   
V. P r i ma A. Syunyakov LN812379LN 812 613. S. caucasica  G.L . Kudryashova LN812381LN812 615.
S. caulescens S. caveana G. & S. Miehe 00-303-12 LN812382LN812 616.
S. cebennensis       N. Tkach 261 & M. Röser 
LN812383LN 81 2617. S. cernua   N.V. Matveeva & L. Zanokha
LN812384LN812618. S. cespitosa cespitosaYu.P. Kozhe vnik ov LN812385LN81261 9. S. cespitosa monticola
V.V. Petrovsky & P.G. Zhukova LN812386LN812620. S. charadzeae T.N. Popova
& G.L. Kudryashova LN812387LN812621. S. cherlerioides  cherlerioidesV.Yu. Bark alov LN812389LN812623.
S. cherlerioides  rebunshirensis N. T kach 252 & M. Röser  LN812388LN812622.
S. chrysantha G.N. Jones LN812390LN812624. S. cintrana S. cochlearis
N. T kach 213 & M. Röser  LN812391LN812625. S. colchica 
A. Dolmatova & al. LN812392LN812626. S. columnaris S. Shkhagansoev LN812393LN812627.
S. conifera       J. Fernandez-Casas 2362 LN812394L N812628.
S. consanguinea G. & S. Miehe & U. Wündisch 94-344-12 LN812395LN812629. S. continentalis    
 A. Segura Zubizarreta LN812396 LN812630. S. contraria rubella   G. & S. Miehe 00-322-02 
LN812397LN812631. S. coriifolia E. Mordak h LN812398LN812632. S. corsica M. Röser
10171  LN812399LN812633. S. cortusifolia E. Pobedimova & G. Konovalova LN812634.
S. corymbosa M. Röser 9855    LN812400 LN812635. S. cossoniana  
S. cotyledon M. Röser 10572  LN812401LN812636. S. cotyledon 
N. Tkach 212 & M. Röser  LN812380L N812 614. S. crustata 
N. Tkach 79  LN812402LN812637. S. cunea ta N. T kach 230 & M. Röser   LN812403
LN812638. S. cuneifolia    N. T kach 71  LN812404LN812639. S. cymbalaria 
T.N. Popova & S.V. Bondarenko LN812405LN812640
S. depressa S. derbekii A.T. Reutt LN812406LN812641. S. desou-
lavyi    N. Tkach 214 & M. Röser   LN812407 LN812642. S. diapensioides 
N. Tkach 231 & M. Röser    LN812408LN812643. S. dichotoma 
Rivas Goday    LN812409LN812644. S. dingqingensis S. dinnikii 
N.N. Portenier LN812410LN8126 45. S. discolor W. Hilbig  LN812411S. diversifolia 
diversifoliaG. & S. Miehe & U. Wündisch 94-510-29 LN812412LN812646. S. diversifolia haematophylla 
S. duthiei  N. Tkach 223 & M. Röser  LN812413L N812647.
Appendix 1.


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TAX ON
–Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
Version of Record
S. egregia S. engleriana G. & S. Miehe 9541/03 LN812 414LN812648. S. erioblasta
M. Röser 3251  LN812415LN812649. S. eschscholtzii P. Zhmy le v E.Yu. Norkina &
T.V. Plieva LN812416LN812650M.H. Hof fmann 11/55 LN812651. S. exarata M. Röser 6547    
LN812417LN812652. S. federici-augusti grisebachii 
N. Tkach 216 & M. Röser  LN812443LN812678. S. felineri   N. Tkach 207 &
M. Röser  LN812419LN81265 4. S. ferdinandi-coburgi N. Tkach 232 & M. Röser
   LN812420LN812655. S. filicaulis Mie he 97-064 -10 Huang Otsu & Tunsu LN812421LN812656. S. finitima
G. Miehe & U. Wündisch 94-118-03  LN812422LN812657. S. firma    T. Derviz-
Sokolovskaya 498 LN812423LN812658Yu.P. Kozhevnikov & M.D. Andreeva LN812424LN812659. S. flagellaris 
crandallii R.E. Brooks 20451 LN812425LN812660. S. flagellaris crassiflagellata 
G. & S. Miehe 1008 LN812426LN8126 61. S. flagellaris stenophylla 
  G. & S . Miehe 6420  LN812427S. flexuosa E. Hultén LN 812 516LN812751. S. florulenta 
S. fortunei fortuneiE. Egorova & L. Kolchanova LN812663. S. fortunei 
incisolobata N. Tkach 215 & M. Röser   LN812428LN812662.
S. fragilis  A. Segura Zubizaretta 38403    LN812429LN812664S. fragosoi
N. T kach 206 & M. Röser  LN812430LN812665. S. funstonii 
A. Taran LN812431LN812666M.P. Andreev E.Yu. Norkina & V.V. Petrovsk y LN812432LN812667. S. gem-
migera gemmuligera G. & S. Miehe & U. Wündisch 94-390-09 LN812433LN812668. S. gem-
mipara S. gemmulosa M. Röser 4738    LN812434LN812669. S. genesiana    
S. georgei   N. Tkach 233 & M. Röser  LN812435LN812670.
S. geranioides M. Röser 10531 LN812436LN812671. S. globulifera M. Röser 4622    
LN812437LN812672V.P. Botschantzev LN812438LN812673. S. gouldii G. & S. Miehe 00 -160-35
LN812439LN8 126 74. S. graeca H.J. Zündorf 23103 LN812440LN812675. S. granulata M. Röser
7437   LN812441LN812676. S. granulifera G. & S. Miehe 00-160-14  LN812442LN812677. S. haenseleri 
S. hariotii S. hederacea 
P. Gutte 62/2001  LN812444LN812679. S. heleonastes   G. & S. Miehe 03-094-26  LN812445LN812680.
S. heterotricha G. Miehe & U. Wündisch 94-150-20 LN812446LN812681. S. hirculoides 
G. & S. Miehe 570 LN812447LN812682. S. hirculoides G. & S . Miehe 99- 046- 09  L N812574
LN812725. S. hirculus hirculus O.M. Afonina  LN812449LN812684M.H. Hof fmann M36  
LN812450LN812685. S. hirculus alpina G. & S. Miehe 5780  LN812448LN812683. S. hirsuta 
S. hispidula G. & S. Miehe 00-205-18 LN812451
LN812686. S. hookeri S. hostii N. Tkach
70    LN812452 LN812687. S. hyperborea A.A. Korobkov LN812453 LN812688. S. hypnoides 
N. Tkach 65   LN812454LN812689. S. implicans G. & S. Miehe 10-27-02 
LN812455LN812690. S. insolens S. intricata S. iranica 
 N. Tkach 234 & M. Röser  LN812456LN812691. S. irrigua J. Wankow 
LN812457LN812692. S. isophylla  G. Miehe & U. Wündisch 94-182-12 LN812458LN812693. S. italica 
 Anonymous  LN812459LN812694. S. jacquemontiana  G. & S. Miehe 00-367-09  LN812460
LN812695. S. juniperifolia S. kingdonii G. Miehe & U. Wündisch 94-173-
10  LN812461 LN812696. S. kingiana   S. kolenatiana  Z. Klochkova
LN812462LN812697. S. komarovii N.N. Tzvelev LN812463LN812698. S. korshinskyi 
S. kotschyi N. T kach 235 & M. Röser  LN812464LN812699. S. kruh-
siana A.A. Korobkov & N.A. Sek retareva LN812465LN812700. S. lactea V.V. P etro vs ky 
LN812466LN812701. S. latepetiolata Anonymous 738    LN812467LN812702
S. latiflora  G. & S. Miehe 00- 421-04 LN812468LN812703. S. lilacina 
N. Tkach 80    LN812469LN812704. S. lingulata M. Röser 6664  LN812470
LN812705. S. litangensis G. Miehe & U. Wündisch 94-136-32 LN812471LN812706. S. luteoviridis 
N. Tkach 236 & M. Röser   LN812472LN812707. S. lychnitis G. & S. Miehe
00 -373 - 04 LN812473LN812708. S. macrocalyx R.V. Kamelin & al. LN812474LN812709. S. maderensis 
K. Werner  LN812475S. magellanica N. Tkach 254 & M. Röser  
LN812476LN812710. S. marginata M. Röser 9695   LN812478LN812712
N. Tkach 229 & M. Röser   LN812363LN812597. S. marginata bubakii 
N. Tkach 217 & M. Röser  LN812477LN812711. S. matta-florida N. Tkach
237 & M. Röser    LN812479LN812713. S. mertensiana S. moncayensis 
 N. Tkach 238 & M. Röser     LN812480LN812714. S. moorcroftiana   G. &
S. Miehe 00-177-02 LN812482LN812716. S. moschata   N. Tkach 82   
LN812483LN812717. S. mucronulata G. Miehe & U. Wündisch 94-251-15 LN812484LN812718. S. mucronulatoides   
G. & S. Miehe 00 -320-04  LN812485LN812719. S. mutata M. Röser 10412 LN812486LN812720. S. nanella
G. & S. Miehe 9388/17 LN812487LN812721. S. nangxianensis S. nevaden-
sis M. Röser 3313 LN812488LN812722. S. nigroglandulifera G. & S. Miehe 98-059-33
LN812723. S. nipponica M. Togasi 1707 LN812489LN812724. S. nishidae 
S. omolojensis A.T. Reutt LN812490LN812726B.A. Jurtzev
LN812491S. oppositifolia S. oppo-
sitifolia   oppositifolia B.A. Jurtzev   LN812727. S. oppositifolia  asiatica    
V.A. Siplivinsk iy 74-106 LN812492LN812728G. & S. Miehe 7705 LN812729. S. oppositifolia small-
iana S. osloensis 
Schwerdtfeger 12431 LN812493LN812730. S. paniculata 
N. Tkach 66  LN812494LN812731. S. paradoxa R. Freiherr von Benz 3095
   LN812495LN812732. S. parnassifolia G. & S. Miehe 00-223-28 LN812496LN812733. S. pedemontana
cymosa M. Röser 9674 LN812497LN812734. S. pentadactylis 
M. Röser 10552   LN812498LN812735A. Segura Zubizarreta LN812499LN812736. S. pentadactylis 
willkommiana S. peplidifolia S. perpusilla 
Appendix 1.



Tkach & al. • Molecular phylogeny of Saxifraga (Saxifragaceae)
TAX ON
–
Version of Record
G. & S. Miehe 00-364-01  LN812500LN812737. S. petraea M. Röser 10236    LN812501LN812738. S. pilifera 
 G. Miehe & U. Wündisch 94-105-28  LN812502LN812739. S. platysepala 
V.V. Pe t r ov s k y LN812503LN812740. S. pontica M. K r…LN812504S. porophylla 
N. Tkach 239 & M. Röser    LN812505LN812741. S. portosanctana S. prae-
termissa M. Röser 10584 LN812506LN812742. S. presolanensis Anonymous 
LN812507LN81274 3. S. przewalskii J.F. Rock 13017 LN81250 8LN81274 4. S. pseudohirculus   G. &
S. Miehe & U. Wündisch 94-332-05   LN812509LN81 2745. S. pseudolaevis Sh. Shetekauri R. Gagnidze & T. Zangu-
rashvili LN812510S. pubescens iratiana N. Tkach 240
& M. Röser     LN812511 LN812746. S. pulvinaria   G. & S. Miehe 505 LN812512LN812747.
S. punctulata S. radiata  E. Hultén  LN812513LN812748 
V.V. P e t rov s k y T.M. Koroleva & M.P. Andreev LN812418LN812653. S. retusa augustana 
N. Tkach 241 & M. Röser    LN812 514LN812749. S. reuteriana S. rigoi 
S. rivularis rivularisV.A. Gavrilyuk & P.K. Gagarin LN812517LN812752. S. riv-
ularis arctolitoralis T.M. Zaslavskaya I.M. Litvinov & T.V. Plieva LN812515
LN812750. S. rosacea M. Röser 1115  LN812518LN812753. S. rotundifolia 
N. Tkach 68  L N812519LN81275 4. S. roylei N. Tk ach 227 & M. Röser
    LN812520LN812755. S. rufescens  A. Favre & S . Matuszak 135 LN812521LN812756. S. ruprechtiana 
  N.N. Tzvelev & al.  LN812757. S. saginoides   G. & S. Miehe 00-361-11 LN812522
LN812758. S. sancta M. Röser 2700   LN812523LN812759. S. sanguinea 
–. S. scardica N. Tkach 242 & M. Röser   LN812524LN812760. S. scleropoda 
  D. Geltman Dolmatova & Shvanova LN812525S. sediformis  J.F. Rock 9833 
LN812526S. sedoides hohenwartii M. Röser 10442  LN812527LN 812761. S. seguieri
M. Röser 1469  LN812528LN812762. S. sempervivum N. T kach
255 & M. Röser   LN812529LN812763. S. serpyllifolia serpyllifolia:  B.A. Jurtzev LN812531 LN812765.
S. serpyllifolia glutinosa Yu. P. Ko zh ev nikov LN812530LN 812764. S. setigera 
S. sibirica M. Mikhailova LN812532LN81276 6. S. sibthorpii J.N.F. Bornmülle r
675 LN812533LN812767. S. sieversiana S. signatella G. Miehe &
U. Wündisch 94-167-11 LN812534 LN812768. S. sinomontana     G. & S. Miehe 00-329-02  LN812480
LN812715G. & S. Miehe & U. Wündisch 94-300-35 LN812535LN812769. S. spathularis 
   M. Röser 9430    LN812536 LN812770. S. spinulosa 
N.V. Matveeva & L . Zanokha LN812537LN812771. S. spruneri   N. Tk ach 243 &
M. Röser  LN812538LN812772. S. squarrosa S. stella-aurea polyadena
 G. & S. Miehe 9436/05 LN812539LN812773. S. stelleriana 
S. stolitzkae N. Tk ach 244 & M. Röser   LN812540LN812774. S. sto-
lonifera  N. Tkach 256 & M. Röser   L N812 541 LN812775. S. stribrnyi 
N. Tkach 218 & M. Röser    LN812542LN812776. S. strigosa ramosa 
S. substrigosa G. Miehe & U. Wündisch 94-177-23 LN812778. S. sub-
verticillata N. Tkach 262 & M. Röser LN812543LN812777.
S. tangutica G. Miehe & U. Wündisch 94-099-20 LN812544LN812779. S. tatsienluensis 
–. S. taygetea L. Martins & J. Müller 1815 LN812545LN812780. S. taylorii 
J.A. Calder D.B.O. Savile & R.L. Taylor 23085 LN812546LN812781. S. tenella 
N. Tkach 69    LN812548 LN812783  N. Tkach 257 & M. Röser    L N8125 47
LN812782. S. terektensis    A. Kuminova N. Peshk ova & S. Linde  LN812549LN812784. S. thessalica 
N. T kach 245 & M. Röser    LN812550LN812785. S. tibetica 
G. & S. Miehe 9378/06  LN812551LN812786. S. tombeanensis N. Tkach
219 & M. Röser LN812552LN812787. S. trabutiana        B. Valdes & al . 60-2033 LN812553
–. S. tricuspidata N. Tkach 205 & M. Röser   LN812554LN812788M.H. Hoff-
mann 11/22   LN812555LN812789. S. tridactylites M. Röser 9927 LN812556LN812790. S. trifurcata
   N. Tkach 220 & M. Röser  LN812557LN812791. S. umbellulata 
umbellulata:G. Miehe & U. Wündisch 94-156-26 LN812559LN812793. S. umbellulata muricola 
G. Miehe & U. Wündisch 94-055-20 LN812558LN812792. S. umbellulata pectinata 
S. umbrosa N. Tkach 72   LN812560
LN812794. S. unguiculata G. & S. Miehe 9497/17   L N81 2561LN812795. S. unguipetala 
 N. Tkach 246 & M. Röser   LN812562LN812796. S. valdensis O. Angerer LN812563
LN812797 S. vandellii N. Tkach 247 & M. Röser
   LN812564LN812798. S. vayredana  N. Tkach 248 & M. Röser
   LN812565LN812799. S. veitchiana N. Tkach 221 & M. Röser    LN812566
LN812800. S. vespertina S. viscidula G. & S. Miehe 00 -262-
01 LN812567LN8128 01. S. wahlenbergii Anonymous LN812568LN812802. S. wallichiana 
G. & S. Miehe 00-228-31 LN812569LN812803. S. wardii G. Miehe & U. Wündisch 94-173-07 LN812570
LN812804. S. wendelboi N. Tkach 222 & M. Röser LN812571LN812805.
S. xiaozhongdianensis S. yarlungzangboensis G. Miehe & U. Wündisch 94-166-06 
LN812572LN812806. S. zhidoensis S. zimmermannii U. Wündisch 278 
LN812573LN812807. Saxifragodes albowiana Sa xifragopsis fra-
garioides Suksdorfia violacea Sullivan-
tia oregana Tan ak a ea radicans 
Telesonix heucheriformis Te lli ma grandiflora 
Tiarella polyphylla        
Tolmiea menziesii  
Appendix 1.
