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Phylogeny of Apiaceae subtribe Daucinae and the taxonomic delineation of its genera

Authors:
  • University of Warsaw Biological and Chemical Research Centre
  • Via Apia, France, Mulhouse

Abstract and Figures

Scandiceae subtribe Daucinae encompasses umbellifers that have fruits with prominent secondary ridges projecting into wings (former tribe Laserpitieae) or spines (former tribe Caucalideae pro parte). It comprises several economically or medicinally important genera including Cuminum, Daucus, Laser, Laserpitium and Thapsia among others. Recent molecular studies, based mostly on nrDNA ITS sequences, revealed that neither Daucus nor Laserpitium are monophyletic. To address issues of relationships and apply respective nomenclatural changes, we obtained additional ITS sequences as well as inde- pendent data from three plastid markers—rps16 intron, rpoC1 intron and rpoB-trnC intergenic spacer—for a comprehensive sample of the subtribe. We examined data for 260 accessions representing all genera of Daucinae and 81 of its ca. 93 species. Phylogenetic trees were estimated using maximum likelihood and Bayesian inference methods. The results indicate that former Laserpitieae constitute a paraphyletic grade at the base of the spiny-fruited members of Daucinae while traditionally delimited Daucus and Laserpitium are polyphyletic. To maintain a monophyletic Daucus, we suggest including the following genera and species into its synonymy: Agrocharis, Melanoselinum, Monizia, Pachyctenium, Pseudorlaya, Rouya, Tornabenea, Athamanta dellacellae and Cryptotaenia elegans. The species of Laserpitium occur in seven clades and only six species of the Laserpitium s.str. clade retain the generic name. Several species are transferred to Ekimia, Laser and Thapsia; additionally, a monospecific genus Siler is restored and a new genus, Silphiodaucus, is established. The inclusion of Ammodaucus into Thapsia suggested in an earlier study is not supported. The position of Laserpitium pseudomeum requires further study.
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Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
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   

SilerElaeose-
linum Margotia 
GuilloneaLaserpitium Melanoselinum
 PolylophiumThapsia Tornabene a 



  
Thapsia
Melanoselinum edule
Phylogeny of Apiaceae subtribe Daucinae and the taxonomic
delineation of its genera
Łukasz Banasiak,1 Aneta Wojewódzka,1 Jakub Baczyński,1 Jean-Pierre Reduron,2 Marcin Piwczyński,3
Renata Kurzyna-Młynik,1 Rafał Gutaker,1 Agnieszka Czarnocka-Cieciura,1 Sylwia Kosmala-Grzechnik1 &
Krzysztof Spalik1
1 Department of Molecular Phylogenetics and Evolution, Institute of Botany, Faculty of Biolog y, University of Warsaw Biological
and Chemical Research Centre, Żwirki i Wigury 4, 02-089 Warszawa, Poland
2 VIA APIA, 10 rue de l’Arsenal, 68100 Mulhouse, France
3 Chair of Ecolog y and Biogeography, Nicolaus Copernicus University in Toruń, Lwowska 1, 87-100 Toruń, Poland
 Krzysztof Spalik, spalik@biol.uw.edu.pl
ORCID 
DOI  
Abstract
 
        
CuminumDaucusLaserLaserpitium and Thapsia
Daucus Laserpitium 
   
r ps16rpoC1rpoB-trnC



Daucus and Laserpitium . Daucus
AgrocharisMelanoselinumMoniziaPachycteniumPseudorlayaRouyaTornabeneaAthamanta
dellacellae and Cryptotaenia elegansLaserpitium Laserpitium
EkimiaLaser and Thapsia
Siler Silphiodaucus Ammodaucus Thapsia 
Laserpitium pseudomeum 
Keywords
Daucus Laserpitium
Supplementary Material



Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
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Tab le . 
 




Agrocharis  A. melanantha  4 4 (3) 
Ammodaucus aA. leucotrichus  2 1 (1) 
Athamanta A. dellacellae  1 1 (1) 
CryptotaeniaC. elegans  1 1 (1) 
Cuminum  C. cyminum  3 2 (2) 
Daucus  D. carota  26 23 (20) 
Ekimia  E. bornmuelleri  1 1 (1) 
Laser  L. trilobum 1 1 (1) 
Laserpitium  L. gallicum  18 (14) c
Melanoselinum  M. decipiens 1 1 (1) 
Monizia  M. edulis 1 1 (1) 
Orlaya O. grandiflora  3 3 (2) 
Pachyctenium  P. mirabile  1 1 (0) 
Polylophium  P. orientale 2 2 (1) 
Pseudorlaya  P. pumila  2 2 (0) 
Rouya  R. polygama  1 1 (1) 
Thapsia dT. villosa 18 15 (8) 

*Distichoselinum D. tenuifolium 1 1 (1) 
*Elaeoselinum E. meoides 4(–5) 3 (2) 

*Guillonea  G. scabra 1 1 (1) 
*Margotia M. laserpitioidesM. gummifera 1 1 (1) 
Tornabenea  T. insularis 6 3 (1) 
a AmmodaucusThapsia 
Thapsia 
Athamanta and Cryptotaenia 
c Laserpitium

dThapsia 

Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
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Monizia 
SilerS. montanum
Laserpitium (a s L. siler 
SilerS. trilobum 
Laser

 Rouya
Thapsia
Tornabe-
neaPolylophium and Thapsia
 
Ekimia    
Laserpitium 
E. bornmuelleri 
Prangos 







Thapsia
Elaeoselinum Margotia Guillonea  Laserpitium
   
  Siler
Laserpitium  
 Tornabenea
 
   

 
Elaeoselinum
Margotia GuilloneaLaser pitiumLaser
Melanoselinum Polylophium Rouya Siler  Thapsia
MoniziaTornabenea
LaserpitiumL. prutenicum and L. his-
pidumDaucus
Silphiodaucus


  

    

   
   
    







rpoC1   
matK


Laserpitium
Daucus

  Thapsia
Laser and Polylophium
MoniziaMelanoselinum


 


Daucus 

Daucus Daucus 
 D. carota 

Melanoselinum and Monizia 
D. carota   Tornabenea
  
DaucusD. carota  
     



Laserpitium


 

  
Daucus



2011). 
 Laser 
LaserpitiumThapsia

    






MATERIALS AND METHODS
Taxon and molecular marker sampling. —





 
Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
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


     



 



rp oC1 and rps16rpoB-trnC



DNA extraction, amplification, and sequencing. —   




  
 


rpoB-trnC


 

  

   
   

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


rps16



 rpoC1  






 
      





  
         

  




Sequence and phylogenetic analyses. —

  




p 
  






 




Tab le . 

rpoB-trnC

rpoC1

rps16
 

 Reduceda
       
  1643    4415 
   111  632 681 
 203 1010 854  2536  
  241  183   
  243 503  3134 3404 
 3.4  1.6 0.4  62.1 
arpoC1rps16 
Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
Version of Record
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
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  



  

RESULTS
Sequence analyses. —
rps16 
rp oC1rpoB-trnC

 
rpoB-trnC

    



  
 
  




   
    
   



  
   



  

  



 r poC1r p s16 




P < 10 
  

Tab le . 

 P

rpoB-trnCrpoC1  
rpoB-trnCrpoC1rps16 
rpoB-trnCrpoC1rps16 *

rpoB-trnC 0.610
rpoB-trnCrpoC1 0.240
rpoB-trnCrpoC1rps16 



Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
568 Version of Record
TAX ON
65 (3) • June 2016: 563 585
Fig. 1.

















Anthriscus sylvestris 83
Glaucosciadium insigne 74
Glaucosciadium cordifolium 221
Ferula communis 195
Ferula olivacea 363
Leutea avicennae 67
Laserpitium latifolium 498
Laserpitium krapfii subsp. gaudinii 497
Laserpitium krapfii subsp. krapfii 496
Laserpitium gallicum subsp. angustissimum 488
Laserpitium gallicum subsp. gallicum 487
Laserpitium gallicum subsp. paradoxum 490
Laserpitium gallicum subsp. orospedanum 489
Ekimia bornmuelleri 655
Laserpitium petrophilum 509
Thapsia scabra 479
Thapsia villosa var. dissecta 528
Laserpitium nestleri subsp. flabellatum 505 Laserpitium nestleri subsp. lainzii 506
Laserpitium nestler i subsp. nestleri 501
Laserpitium eliasii subsp. thalictrifolium 486
Laserpitium eliasii subsp. eliasii 484
Laserpitium eliasii subsp. ordunae 485
Thapsia meoides 473
Thapsia gummifera 520
Thapsia sp. 471
Thapsia thapsioides 478
Thapsia thapsioides 477
Thapsia tenuifolia 469
Thapsia transtagana 524
Thapsia garganica 527
Thapsia garganica 526
Thapsia garganica 525
Cuminum cyminum G7
Cuminum setifolium 20
Cuminum cyminum 339
Laser trilobum 611
Laser trilobum G61
Laserpitium stevenii 518
Ammodaucus leucotrichus 12
Laserpitium carduchorum 483
Polylophium panjutinii G82
Laserpitium affine 480
Laserpitium archangelica 481
Laserpitium pseudomeum 513
Laserpitium siler 515
Laserpitium siler 516
Laserpitium siler G70
Laserpitium siler 517
Orlaya grandiflora 258
Orlaya daucoides G75
Laserpitium pr utenicum subsp. prutenicum 649
Laserpitium hispidum 494
Agrocharis incognita 245
Agrocharis pedunculata 247
Agrocharis incognita 46
Daucus broteri 276
Daucus broteri 277
Daucus broteri 440
Daucus conchitae 236
Daucus involucratus 37
Daucus involucratus 460 272
Daucus littoralis 8
Daucus bicolor 41
Daucus littoralis 461
Daucus durieua 40
Daucus durieua 453
Daucus durieua G38
Daucus arcanus 260
Daucus pusillus G52
Daucus pusillus 466 G53
Daucus guttatus 238 458
Daucus glochidiatus 455
Daucus glochidiatus 456
Daucus montanus 17
Monizia edulis 13 G74
Melanoselinum decipiens G72 G73
Cryptotaenia elegans 39
Daucus tenuisectus 266
Daucus muricatus 281
Daucus muricatus G50
Daucus muricatus G51
Daucus muricatus 271
Daucus crinitus 14
Daucus setifolius 467
Daucus setifolius 237 G56
Daucus aureus G18
Daucus mauritii G45
Athamanta dellacellae 435
Rouya polygama 523
Daucus syrticus 468
Tornabenea annua 521
Daucus carota subsp. gummifer 445 Daucus carota subsp. hispanicus G30
Daucus carota subsp. gadecaei 34
Daucus carota subsp. azoricus 610 Daucus carota subsp. halophilus 9
Daucus virgatus 659
Daucus carota subsp. carota G98
Daucus carota subsp. carota 32 443 444 Daucus carota subsp. commutatus 3
Daucus carota subsp. maritimus 448
0.01
94/1.0
72/0.99
100/1.0
96/1.0
94/1.0
60/−
66/0.56
49/0.50
100/1.0
94/0.98
100/1.0
33/−
28/
100/1.0
82/1.0
97/1.0
98/1.0
95/1.0
100/1.0
100/1.0
81/0.50
87/1.0
88/1.0
52/0.69
100/1.0
88/0.98
95/1.0
100/1.0
27/−
13/−
100/1.0
100/1.0
47/−
100/1.0
13/−
26/−
29/−
55/0.65
96/1.0
27/−
33/−
100/1.0
90/1.0
100/1.0
100/1.0
100/1.0
100/1.0
97/1.0
100/1.0
72/0.95
100/1.0
93/1.0
100/1.0
71/0.94
99/1.0
98/1.0
99/1.0
44/−
97/1.0
100/1.0
31/−
100/1.0
33/−
95/1.0
88/0.81
99/1.0
95/0.99
61/0.69
54/0.78
60/0.90
100/1.0
64/0.63
52/0.94
37/0.73
100/1.0
98/1.0
100/1.0
99/1.0
97/1.0
100/1.0
100/1.0
61/0.94
Laserpitium s.str.
Daucus I
Macaronesian
Daucus II
Agrocharis
Silphiodaucus
Orlaya
Siler
Laser
Cuminum
Thapsia
Ekimia
Outgroup
Daucus s.l.
Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
Version of Record
TAX ON
65 (3) • June 2016: 563 585
Fig. .

rps16 and r po C1
rpoB-trnC













Leutea avicennae 67
Ferula olivacea 363
Ferula communis 195
Anthriscus sylvestris 83
Glaucosciadium insigne 74
Glaucosciadium cordifolium 221
Cuminum cyminum G7
Cuminum setifolium 20
Cuminum cyminum 339
Laserpitium krapfii subsp. krapfii 496
Ammodaucus leucotrichus 12
Laserpitium petrophilum 509
Laserpitium latifolium 498
Laserpitium krapfii subsp. gaudinii 497
Laserpitium gallicum subsp. orospedanum 489
Ekimia bornmuelleri 655
Laserpitium gallicum subsp. angustissimum 488
Laserpitium gallicum subsp. paradoxum 490
Laserpitium gallicum subsp. gallicum 487
Laserpitium nestler i subsp. nestleri 501
Laserpitium eliasii subsp. eliasii 484
Laserpitium eliasii subsp. thalictrifolium 486
Laserpitium nestler i subsp. flabellatum 505
Laserpitium nestleri subsp. lainzii 506
Laserpitium eliasii subsp. ordunae 485
Thapsia scabra 479
Thapsia villosa var. dissecta 528
Thapsia transtagana 524
Thapsia garganica 527
Thapsia garganica 526
Thapsia garganica 525
Thapsia gummifera 520
Thapsia tenuifolia 469
Thapsia sp. 471
Thapsia thapsioides 478
Thapsia meoides 473
Thapsia thapsioides 477
Laserpitium siler G70
Laserpitium siler 517
Laserpitium siler 516
Laserpitium siler 515
Orlaya daucoides G75
Orlaya grandiflora 258
Laserpitium affine 480
Laserpitium carduchorum 483
Laser trilobum 611
Laserpitium archangelica 481
Laserpitium stevenii 518
Laser trilobum G61
Polylophium panjutinii G82
Laserpitium pseudomeum 513
Laserpitium hispidum 494
Laserpitium pr utenicum subsp. prutenicum 649
Agrocharis incognita 245
Agrocharis pedunculata 247
Agrocharis incognita 46
Daucus montanus 17
Daucus glochidiatus 456
Daucus glochidiatus 455
Daucus durieua 40
Daucus durieua 453
Daucus durieua G38
Daucus guttatus 458
Daucus guttatus 238
Daucus pusillus 466
Daucus pusillus G52
Daucus arcanus 260
Daucus pusillus G53
Daucus littoralis 8
Daucus bicolor 41
Daucus littoralis 461
Daucus involucratus 37
Daucus involucratus 460
Daucus involucratus 272
Daucus broteri 276
Daucus conchitae 236
Daucus broteri 277
Daucus broteri 440
Cryptotaenia elegans 39
Melanoselinum decipiens G73
Melanoselinum decipiens G72
Monizia edulis 13
Monizia edulis G74
Daucus muricatus 271
Daucus muricatus 281
Daucus muricatus G51 G50
Daucus mauritii G45
Daucus aureus G18
Athamanta dellacellae 435
Daucus setifolius 237
Daucus crinitus 14
Daucus setifolius 467
Daucus setifolius G56
Daucus tenuisectus 266
Rouya polygama 523
Daucus carota subsp. hispanicus G30
Daucus carota subsp. maritimus 448
Daucus carota subsp. gummifer 445
Daucus carota subsp. azoricus 610 Daucus carota subsp. carota 443 444
Daucus carota subsp. commutatus 3
Daucus carota subsp. carota 32 Daucus carota subsp. halophilus 9
Tornabenea annua 521
Daucus carota subsp. carota G98
Daucus virgatus 659
Daucus carota subsp. gadecaei 34
Daucus syrticus 468
0.001
94/0.99
90/0.99
79/1.0
100/1.0
100/1.0
94/0.99
99/1.0
100/1.0
80/0.96
37/0.50
23/−
24/0.68
74/1.0
96/1.0
40/0.68
61/0.82
100/1.0
86/1.0
91/1.0
34/−
93/1.0
75/0.92
73/0.97
64/0.97
100/1.0
93/1.0
99/1.0
88/1.0
88/1.0
59/0.98
61/0.99
95/1.0
17/0.90
62/0.99
37/0.67
49/0.73
21/0.95
64/0.67
99/1.0
93/1.0
85/1.0
94/1.0
50/0.70
96/1.0
94/0.98
26/0.88
92/1.0
88/0.99
85/1.0
97/1.0
100/1.0
79/1.0
41/0.82
98/1.0
100/1.0
44/0.96
64/0.88
86/1.0
45/0.57 58/−
42/0.96
88/1.0
18/0.52
53/0.97
49/0.97
72/1.0
70/0.95
95/1.0
6/−
99/1.0
32/0.92
93/0.84
62/0.53
79/−
94/0.58
54/0.69
89/0.85
95/0.97
52/−
Laserpitium s.str.
Daucus I
Macaronesian
Daucus II
Agrocharis
Silphiodaucus
Orlaya
Siler
Laser
Cuminum
Thapsia
Outgroup
Daucus s.l.
Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
 Version of Record
TAX ON
65 (3) • June 2016: 563 585
  

P 
    

      
   


   



Phylogenetic analyses of separate nrDNA and cpDNA data.


  






Ekimia



Laserpitium L. gallicum 
 Laserpitium
   

  
L. gal-
licum 

   
L. krapfii krapfii 
 
   


Ekimia bornmuelleri and Laser-
pitium petrophilum 
Ekimia
  



  Thapsia    
Distichoselinum Elaeoselinum
GuilloneaMargotiaLaserpitiumL. eliasii
L. nestleri  Thapsia
  Ammodaucus leucotrichus 


Thapsia 
Thapsia

Thapsia 




Ammodaucus leucotrichus Thapsia



Cuminum cyminum C. setifolium

  


 
  Cuminum
Cuminum
Laser

Laser Laser
   Polylophium and Laserpitium  


Ammodaucus leucotrichus
Ammodaucus  


Laserpitium siler    


Siler
  Siler 
Laserpitium pseudomeum  

Orlaya 

 

Agrocharis Daucus.

 Daucus s.l. clade and
Silphiodaucus    
AgrocharisDaucus 
Daucus 

Silphiodaucus Laserpitium
L. prutenicum and L. hispidum 
Daucus  Silphiodaucus 
Daucus s.l. clade

 Daucus
Daucus 

Daucus 

 
Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
Version of Record
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65 (3) • June 2016: 563 585
Daucus 
Monizia and Melanoselinum
  Cryptotaenia elegans 

Daucus
Tornabenea
and RouyaTornabenea
D. carota

Athamanta dellacellae
 Daucus  

Agrocharis Daucus

Phylogenetic analyses of combined data. —
       
 



rpoC1 
rps16  

  




 

Cuminum 

 
Ammodaucus 
   Thapsia 
Laser  
Ekimia 
Laserpitium glaucum 

Laserpitium  

L. gallicum L. halleri 
L. krapfii    
gaudinii L. nitidum 
L. peucedanoides

Thapsia 
Thapsia 

L. nestleri and L. eliasii    
Thapsia  

Elaeoselinum asclepium  meoides
 Thapsia meoides 


LaserpitiumL. siler and L. pseudomeum

Silphiodaucus 
Daucus s.l. clade.
  
Daucus   
  
Cryptotaenia elegans 
Daucus 
Daucus 


D. carota To r n a -
benea and Daucus virgatus 
Rouya polygama  
D. carota. Daucus 
Pseudorlaya Atha-
manta dellacellae Pachyctenium mirabile 
Pachyctenium  . 
Daucus   
D. broteri D. bicolor 
D. conchitae 
D. involucratus 

Morphological analyses. —
 
        
 



  


  
Daucus
s.l. clade  Silphiodaucus clade and Orlaya) 

OrlayaDaucus  Daucus 
Agrocharis 
Silphiodaucus


Daucus 
DISCUSSION
The incongruence between plastid and nuclear data.

   
        
 

    
 
   


Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
 Version of Record
TAX ON
65 (3) • June 2016: 563 585
Leutea avicennae 67
Ferula communis 195
Ferula olivacea 363 Glaucosciadium cordifolium 221
Glaucosciadium insigne 74 Anthriscus sylvestris 83
Conopodium arvense 630
Athamanta cretensis 617
Cuminum cyminum G7
Cuminum setifolium G6
Cuminum setifolium 20
Cuminum cyminum 339
Cuminum cyminum G10
Cuminum cyminum G12
Cuminum cyminum G11 G13 G8
Cuminum cyminum G9 Ammodaucus leucotrichus G114
Ammodaucus leucotrichus 12
Ekimia bornmuelleri 655
Laserpitium glaucum 491
Laserpitium petrophilum 509
Laserpitium petrophilum G67
Laserpitium petrophilum 508 G85
Laserpitium petrophilum G86
Laserpitium petrophilum G88
Laserpitium petrophilum G87
Laserpitium haller i 493
Laserpitium gallicum subsp. orospedanum 489
Laserpitium gallicum subsp. angustissimum 488
Laserpitium gallicum subsp. paradoxum 490
Laserpitium gallicum subsp. gallicum 487
Laserpitium krapfii subsp. krapfii 496
Laserpitium latifolium 498
Laserpitium krapfii subsp. gaudinii 497
Laserpitium peucedanoides 510
Laserpitium nitidum 507
Laserpitium eliasii subsp. thalictrifolium 486
Laserpitium eliasii subsp. eliasii 484
Laserpitium eliasii subsp. ordunae 485
Laserpitium nestleri subsp. nestleri 501
Laserpitium nestleri subsp. lainzii 506
Laserpitium nestleri subsp. flabellatum 505
Thapsia scabra 479 Thapsia maxima G122 G123
Thapsia villosa G95
Thapsia villosa var. dissecta 528
Thapsia laciniata G132 G133
Thapsia villosa var. dissecta 529
Thapsia villosa G124
Thapsia villosa G125
Thapsia villosa 530 G127
Thapsia villosa G126
Thapsia minor G130
Thapsia minor G131
Thapsia minor G128
Thapsia minor G129
Thapsia asclepium G121
Thapsia asclepium 470
Thapsia tenuifolia 469
Thapsia gummifera 520
Thapsia sp. 471
Thapsia thapsioides 478
Thapsia thapsioides 477
Thapsia meoides 472
Thapsia meoides 473
Thapsia meoides G59
Thapsia smittii G119
Thapsia smittii G120
Thapsia transtagana 524
Thapsia transtagana G116
Thapsia transtagana G94
Thapsia transtagana G117
Thapsia platycarpa G93
Thapsia gymnesica G115
Thapsia garganica 527
Thapsia garganica 526
Thapsia garganica G92
Thapsia garganica 525
Thapsia garganica G128
Laserpitium siler G70
Laserpitium siler G113
Laserpitium siler 517
Laserpitium siler 516
Laserpitium siler 515
Laserpitium pseudomeum 513 Laserpitium affine 480
Laserpitium affine G62
Laserpitium carduchor um 483
Laserpitium carduchor um 482
Laser trilobum G61
Laser trilobum G112
Laser trilobum 611
Laserpitium archangelica 481
Laserpitium stevenii G71
Laserpitium stevenii 518
Polylophium involucratum 395 G81
Polylophium panjutinii G82
Polylophium panjutinii G83
Polylophium panjutinii G84Orlaya grandiflora G77
Orlaya grandiflora 259 G78
Orlaya grandiflora 258
Orlaya daucoides 263
Orlaya daucorlaya G76
Orlaya daucoides G75
Orlaya daucoides 229
Orlaya daucoides G111
Orlaya daucoides 264
Orlaya daucoides G79
Laserpitium prutenicum subsp. duforianum 511
Laserpitium prutenicum subsp. prutenicum 649
Laserpitium pr utenicum subsp. prutenicum G69
Laserpitium hispidum 494
Laserpitium hispidum G64
Laserpitium hispidum 495
Laserpitium hispidum G66
Laserpitium G65
Daucus s.l. clade
0.001
100/1.0
99/1.0
84/1.0
97/1.0
100/1.0
94/1.0 96/1.0
100/1.0
100/1.0
96/0.99
74/0.97
63/0.99
41/−
100/1.0
8/−
100/1.0
96/1.0
100/1.0
72/0.86
9/−
48/0.71
85/1.0
96/1.0
96/1.0
99/1.0 100/1.0
92/1.0
73/0.95
97/1.0
95/0.94
98/1.0
92/1.0
100/1.0
99/1.0
99/1.0
96/0.99
99/1.0
100/1.0
100/1.0
99/1.0
66/0.91
69/0.94
77/0.98
89/1.0
90/1.0
100/1.0
82/0.99
87/1.0
84/1.0
85/0.87
52/0.52
89/1.0
96/1.0
82/0.92
100/1.0
100/1.0
100/1.0
100/1.0
94/−
49/0.80
100/1.0
32/−
45/−
55/0.97
35/0.52
99/1.0
59/1.0
100/1.0
100/1.0
55/1.0
98/1.0
100/0.98
34/0.67
87/1.0
100/1.0
100/1.0
99/1.0
72/0.99
100/1.0
100/1.0
99/1.0
87/0.93
98/1.0
72/0.99
100/1.0
100/1.0
95/0.95
100/1.0
99/1.0
98/1.0
86/1.0 hispidum
Laserpitium s.str.
Silphiodaucus
Orlaya
Siler
Laser
Cuminum
Thapsia
Ekimia
Outgroup
Fig. 3.




Daucus s.l.







Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
Version of Record
TAX ON
65 (3) • June 2016: 563 585
Fig. 4.
Daucus s.l. clade





Daucus carota
maximusDaucus
virgatus 50.
Agrocharis gracilis G1
Agrocharis melanantha G2
Agrocharis melanantha 248
Agrocharis incognita 245
Agrocharis pedunculata G4
Agrocharis pedunculata G3
Agrocharis pedunculata 247
Agrocharis incognita 46 Daucus durieua 40
Daucus durieua 454
Daucus durieua G36
Daucus durieua G37
Daucus durieua 453
Daucus durieua G38
Daucus glochidiatus G39
Daucus glochidiatus G40
Daucus glochidiatus 455
Daucus glochidiatus 456
Daucus montanus 17
Daucus montanus G47
Daucus montanus 230
Daucus montanus 262
Daucus broteri 276
Daucus broteri 277
Daucus broteri 440
Daucus conchitae G32
Daucus conchitae 236
Daucus involucratus 37
Daucus involucratus 460
Daucus involucratus G44
Daucus involucratus 272
Daucus pusillus 466
Daucus pusillus G52
Daucus pusillus G53
Daucus pusillus 267
Daucus arcanus 260
Daucus arcanus G14
Daucus arcanus G15
Daucus guttatus G43
Daucus guttatus 238
Daucus guttatus 458
Daucus littoralis 275
Daucus littoralis 8
Daucus littoralis 461
Daucus bicolor 41
Daucus bicolor G42
Monizia edulis G110
Monizia edulis 13
Monizia edulis G74
Melanoselinum decipiens G73
Melanoselinum
Melanoselinum decipiens 612
Melanoselinum decipiens G72
Melanoselinum decipiens 522 613 Cryptotaenia elegans 39
Cryptotaenia elegans G105 G99
Daucus tenuisectus G58
Daucus tenuisectus 36
Daucus tenuisectus 266
Daucus muricatus G48
Daucus muricatus 271
Daucus muricatus 281
Daucus muricatus G51
Daucus muricatus G50
Daucus muricatus G49
Daucus crinitus G33 G34
Daucus crinitus 14
Daucus setifolius 467
Daucus setifolius G55
Daucus setifolius G56
Daucus setifolius 237
Daucus mauritii G45
Daucus mauritii G46 Daucus aureus G17
Daucus aureus G16
Daucus aureus G19
Daucus aureus G18
Daucus aureus 273 439
Athamanta dellacellae 435
Pachyctenium mirabile G80
Pachyctenium mirabile 244
Pseudorlaya pumila 228 G91
Pseudorlaya pumila 227
Pseudorlaya pumila G108
Pseudorlaya minuscula G90
Daucus biseriatus G89
Rouya polygama 523
Rouya polygama G107 Daucus sahariensis G54
Daucus gracilis 270
Daucus gracilis G41
Daucus syrticus G57
Daucus syrticus 468
Daucus syrticus 265
Daucus carota subsp. halophilus 9
Daucus carota subsp. azoricus 610
Daucus carota subsp. carota G22
Daucus carota subsp. halophilus G29
Daucus carota subsp. azoricus G24 Daucus carota subsp. gadecaei G27
Daucus carota subsp. gummifer G28
Daucus carota subsp. gummifer 445
Tornabenea tenuissima G97
Tornabenea annua 521
Tornabenea annua G96
Daucus carota subsp. gadecaei 34
Tornabenea insularis G106
Daucus carota subsp. drepanensis G26
Daucus carota subsp. hispanicus 447
Daucus carota subsp. hispanicus 35
Daucus carota subsp. hispanicus G30
Daucus carota subsp. carota 32
Daucus virgatus 659
Daucus carota subsp. carota 442 Daucus carota subsp. maximus 274
Daucus carota subsp. carota G25
Daucus carota subsp. maximus 451
Daucus carota subsp. carota G98
Daucus carota subsp. maritimus 448
Daucus carota subsp. carota 443 444
Daucus carota subsp. maximus 450
Daucus carota subsp. hispanicus var. linearis 38 *
Daucus carota subsp. maximus G31
Daucus carota subsp. capillifolius G21
Daucus carota subsp. carota G23
Daucus carota subsp. commutatus 3
0.001
100/1.0
100/1.0
100/1.0
89/1.0
55/0.83 95/1.0
51/0.70
100/1.0
64/0.90
100/1.0
98/1.0
100/1.0
95/1.0
96/1.0
90/0.58
95/1.0
64/0.98
64/0.96
99/1.0
100/1.0
90/1.0
99/1.0
72/0.98
79/1.0
95/1.0
49/0.69
100/1.0
63/0.85
93/0.99
46/0.62
100/1.0
99/0.97
100/1.0
100/1.0
99/1.0 100/1.0
100/1.0
96/1.0
97/1.0
95/1.0
30/−
67/0.74
100/1.0
86/0.96
70/0.96
100/1.0
100/1.0
63/0.76
65/0.98
79/1.0
95/1.0
100/1.0
100/1.0
93/0.92
87/1.0
100/1.0
100/1.0
100/1.0
87/0.98
86/1.0
82/1.0
85/0.99
100/1.0
86/1.0
100/1.0
83/0.99
81/0.91
56/0.86
100/1.0
52/0.88
58/0.92
100/1.0
99/1.0
100/1.0
94/0.96
100/1.0
62/0.91
65/0.98 84/0.98
18/−
57/0.92
3/−
65/0.74
1/−
Daucus I
Macaronesian
Daucus II
Agrocharis
decipiens G109
Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
 Version of Record
TAX ON
65 (3) • June 2016: 563 585
Fig. .


Laserpitium s.str.
Daucus I
Macaronesian
Daucus II
Agrocharis
Silphiodaucus
Orlaya
Laser
Cuminum
Thapsia
Ekimia
Outgroup
Daucus s.l.
Leutea avicennae
Ferula communis
Ferula olivacea
Glaucosciadium cordifolium
Glaucosciadium insigne
Anthriscus sylvestris
Conopodium arvense
Athamanta cretensis
Cuminum cyminum
Cuminum setifolium
Ammodaucus leucotrichus
Ekimia bornmuelleri
Laserpitium petrophilum
Laserpitium glaucum
Laserpitium halleri
Laserpitium gallicum subsp. orospedanum
Laserpitium gallicum subsp. angustissimum
Laserpitium gallicum subsp. paradoxum
Laserpitium gallicum subsp. gallicum
Laserpitium krapfii subsp. krapfii
Laserpitium latifolium
Laserpitium krapfii subsp. gaudinii
Laserpitium peucedanoides
Laserpitium nitidum
Thapsia asclepium
Thapsia garganica
Thapsia gymnesica
Thapsia transtagana
Thapsia platycarpa
Thapsia tenuifolia
Thapsia gummifera
Thapsia thapsioides
Thapsia smittii
Thapsia meoides
Thapsia scabra
Thapsia maxima
Thapsia villosa
Thapsia minor
Thapsia villosa var. dissecta
Thapsia laciniata
Laserpitium eliasii subsp. thalictrifolium
Laserpitium eliasii subsp. eliasii
Laserpitium eliasii subsp. ordunae
Laserpitium nestleri subsp. nestleri
Laserpitium nestleri subsp. lainzii
Laserpitium nestleri subsp. flabellatum
Laserpitium siler
Laserpitium pseudomeum
Laserpitium affine
Laserpitium carduchorum
Laser trilobum
Laserpitium archangelica
Laserpitium stevenii
Polylophium panjutinii
Polylophium involucratum
Orlaya grandiflora
Orlaya daucoides
Orlaya daucorlaya
Laserpitium hispidum
Laserpitium prutenicum subsp. prutenicum
Laserpitium prutenicum subsp. duforianum
Agrocharis pedunculata
Agrocharis incognita
Agrocharis gracilis
Agrocharis melanantha
Daucus durieua
Daucus montanus
Daucus glochidiatus
Daucus broteri
Daucus involucratus
Daucus conchitae
Daucus arcanus
Daucus pusillus
Daucus guttatus
Daucus bicolor
Daucus littoralis
Melanoselinum decipiens
Monizia edulis
Cryptotaenia elegans
Daucus muricatus
Daucus tenuisectus
Daucus setifolius
Daucus crinitus
Daucus aureus
Daucus mauritii
Pachyctenium mirabile
Athamanta della−cellae
Pseudorlaya pumila
Pseudorlaya minuscula
Daucus biseriatus
Rouya polygama
Daucus gracilis
Daucus sahariensis
Daucus syr ticus
Daucus carota subsp. gummifer
Daucus carota subsp. gadecaei
Daucus carota subsp. halophilus
Daucus carota subsp. azoricus
Tornabenea annua
Tornabenea tenuissima
Tornabenea insularis
Daucus carota subsp. drepanensis
Daucus carota subsp. hispanicus
Daucus virgatus
Daucus carota subsp. maximus
Daucus carota subsp. capillifolius
Daucus carota subsp. commutatus
Daucus carota subsp. carota
Daucus carota subsp. maritimus
Morphology of secondary ribs
obsolete
winged
spiny
keeled
Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
Version of Record
TAX ON
65 (3) • June 2016: 563 585


   
  



Thapsia 

T. villosa   
Daucus 
D. glochidiatus 
D. montanus


The homoplasy of fruits with winged secondary ribs. —
 





  


  
  

 
 
    


 
     
   
 




Daucus and Laserpi-
tium

The phylogeny and ta xonomy of Daucus. —
 
Daucus carota sativus
Daucus 


Chamaepsila rosae
 
D. carota capillifolius
. 
D. carota 


Daucus
  Daucus 
Daucus   


Daucus   
Pachyctenium and PseudorlayaDaucus 
Agrocharis
Athamanta dellacellae and
TornabeneaDaucus 

Daucus 


Agrocharis
Daucus 

  Daucus.
Daucus 
 



 
Daucus 




 
Daucus

Agrocharis and Pseudorlaya
MelanoselinumMonizia Pachyctenium Rouya and Tornabe-
nea Daucus.
Daucus s.l.
Daucus 
Daucus.
Daucus 
DaucusPlatyspermum     
Anisactis  
Plat yspermum 
D. muricatus 
DaucusPlatysper mum
DaucusAnisactis
D. brachiatus
D. glochidiatusD. toriloides D. mon-
tanusD. montevidensis  D. pusillus
and D. australis  D. montanus 
Daucus 
Daucus Anisactis
D. brachiatus
 Daucus
Agrocharis and Melanoselinum
Daucus.
Rouya. — Rouya poly-
gamaThapsia

Banasiak & al. • Phylogeny of Apiaceae subtribe Daucinae
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65 (3) • June 2016: 563 585



Daucus 
   
Daucus
polygamus
Daucus D. polygamus 
  
D. carota drepanensis
   
D. bicolor
rouyi 


Flore de France.
Pseudorlaya. —
 P. pumila P. minuscula 
 

Daucus 
