The borage family (Boraginaceae s.str.): A revised infrafamilial classification based on new phylogenetic evidence, with emphasis on the placement of some enigmatic genera

Abstract

Boraginaceae s.str. is a subcosmopolitan family of 1600 to 1700 species in around 90 genera, and recent phylogenetic studies indicate that the infrafamilial classification as currently used is highly obsolete. The present study addresses the relationships of the major clades in Boraginaceae s.str. with an emphasis on monophyly of, and relationships between previously recognized clades and the position of various unplaced genera such as Afrotysonia, Anoplocaryum, Brachybotrys, Chionocharis, Craniospermum, Thyrocarpus, and Trigonocaryum using three plastid markers and a taxon sampling with four outgroup and 170 ingroup species from 73 genera. The phylogeny shows high statistical support for most nodes on the backbone and within individual clades. Echiochileae are confirmed as sister to the remainder of Boraginaceae s.str., which, in turn, fall into two well-supported clades, the Boragineae + Lithospermeae and the Cynoglosseae s.l. The latter is highly resolved and includes the Lasiocaryum-clade (Chionocharis, Lasiocaryum, Microcaryum) and the Trichodesmeae (Caccinia, Trichodesma) as sister to the remainder of the group. Rochelieae (formerly the Eritrichieae s.str., also including Eritrichium, Hackelia, and Lappula) form a poorly supported polytomy together with the Mertensia-clade (also including Anoplocaryum, Asperugo, and Memoremea) and the Omphalodes-clade. The enigmatic genus Craniospermum (Craniospermeae) is sister to an expanded Myosotideae (also including Brachybotrys, Decalepidanthus, Trigonocaryum, and Trigonotis) and these two clades are in turn sister to the Core-Cynoglosseae, in which Afrotysonia glochidiata and Thyrocarpus sampsonii are included. Core-Cynoglosseae again fall into two pairs of well-supported subclades. The majority of generic placements are now resolved satisfactorily and the remaining phylogenetic questions can be clearly delimited. Based on the extensive phylogenetic data now available we propose a new infrafamilial classification into three subfamilies and 11 tribes, representing a consensus among the participating authors, according to which major clades are renamed.

Full text

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
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65 (3) • June 2016: 523–546
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INTRODUCTION
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     
The borage family (Boraginaceae s.str.): A revised infrafamilial
classification based on new phylogenetic evidence, with emphasis
on the placement of some enigmatic genera
Juliana Chacón, Federico Luebert,, Hartmut H. Hilger, Svetlana Ovchinnikova, Federico Selvi,
Lorenzo Cecchi, C. Matt Guilliams, Kristen Hasenstab-Lehman, Karel Sutorý, Michael G. Simpson &
Maximilian Weigend
1 Universität Bonn, Nees-Institut für Biodiversität der Pflanzen, Meckenheimer Allee 170, 53115 Bonn, Germany
2 Universidad de Chile, Departamento de Silvicultura y Conservación de la Naturaleza, Santiago, Chile
3
Freie Universität Berlin, Institut für Biologie-Botanik, Altensteinstr. 6, 14195 Berlin, Germany
4 Central Siberian Botanical Garden, SB RAS, Zolotodolinskaya str. 101, 630090 Novosibirsk, Russia
5 Università di Firenze, Dipartimento di Scienze delle Produzioni Agroalimentari e dell’Ambiente (DISPAA), Laboratorio di
Botanica, P. le delle Cascine 28, 50144 Florence, Italy
6 Università di Firenze, Museo di Storia Naturale, sezione botanica “Filippo Parlatore”, via G. La Pira 4, 50121 Florence, Italy
7 Santa Barbara Botanic Garden, 1212 Mission Canyon Road, Santa Barbara, California, 93105, U.S.A.
8 Rancho Santa Ana Botanic Garden, 1500 N College Ave, Claremont, California, 91711, U.S.A.
 Department of Botany, Moravian Museum, Hviezdoslavova 29a, 627 00 Brno, Czech Republic
10 Department of Biology, San Diego State University, San Diego, California 92182, U.S.A.
 Maximilian Weigend, mweigend@uni-bonn.de
ORCID
 
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Abstract
167 0 
 
  
AfrotysoniaAnoplocaryumBrachybotrysChionocharis
Craniospermum ThyrocarpusTrigonocaryum  


Lasiocaryum
(Chionocharis Lasiocaryum MicrocaryumCaccinia Trichodesma) 
.Eritrichium Hackelia and Lappula
MertensiaAnoplocaryum Asperugo and Memoremea Omphalodes
Craniospermum      Brachybotrys
Decalepidanthus Trigonocaryum Tr igo notis  
Afrotysonia glochidiata and Thyrocarpus sampsonii 



Keywords

Supplementary Material



Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
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65 (3) • June 2016: 523–546
 
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
 
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
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


  
   
Omphalodes
Cynoglossum 



Omphalodes
 Memoremea
Nihon  
OmphalodesCynoglossum


included in Cynoglossum



  




       
Afrotysonia  Anoplocaryum Craniosper-
mum Thyrocarpus Afrotysonia


 Tysonia 
 Caccinia  Solenan-
thus  Rindera  

Myosotidium 

Afrotysonia 
Paracynoglossum   
 

Afrotysonia
 Lindelofia Mattiastrum
Paracaryum Trachelanthus 
Dasynotus
  
Afrotysonia 


Anoplocaryum 


Anoplocaryum and Mer-
tensia 




Tr igonot i s  and Asperugo 
Craniospermum







Craniospermum
Trichodesma 
 
Fig. .A,Moritzia ciliata B,
(Onosma alborosea  M. Weigend 9450
C & D,
Lasiocaryum 
C,
Chionocharis hookeri 
D. Boufford 31100D, Lasiocaryum trichocarpum D. Boufford 42157E & F,E, Tri cho -
desma zeylanicum   F, Caccinia macranthera 
O. Mohr 592G–I, MertensiaG, Anoplocaryum compressumH.H. Hilger 1609H, Asperugo
procumbens
I,
Mertensia bakeri M. Weigend 9344
J & K,
Omphalodes
J,
Myosotidium
hortensia  M. Weigend 9086K, Omphalodes verna L,
(Eritrichium aretioides M. Weigend 9126M & N,M, Craniospermum mongolicum 
N, Craniospermum subvillosum 
O & P,O, Decalepidanthus primuloides 
M. Ackermann s.n.P, Myosotis alpestris Q & R, CryptanthaQ, Amsinckia vernicosa 
M. Weigend 9008R, Cryptantha virginensis S & T, Cynoglossum S, Solenanthus
apenninusWeigend 8139T, Trachelanthus cerinthoides 


Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
525Version of Record
TAX ON
65 (3) • June 2016: 523–546

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
526 Version of Record
TAX ON
65 (3) • June 2016: 523–546
Caccinia


Thyrocarpus  

Both-
riospermum 


  

Brachybotrys Chi
-
onocharisTrigonocaryum 



Brachybotrys paridiformis 


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  
  
Trigon otisCerinthe Mertensia
Brachybotrys 



Cerinthe and Mertensia  


Chionocharis hookeri


.
Lasiocaryum

   

LasiocaryumMicrocaryum


Trigonocaryum
Myosotis involucrata

Anchusa 


  



 Myosotis      
Trigonocaryum involucratum 


       
 
  
. 



AfrotysoniaAnoplocaryum
CraniospermumThyrocarpus
BrachybotrysChionocharis
and Trigonocaryum

trnL-trnFrps16
trnS-G 



 

MATERIALS AND METHODS
Taxon sampling. —      
    


Afrotysonia 
A. glochidiata  Anoplocaryum 
A. compressumA. hele-
nae Craniospermum C. echioides
C. mongolicumC. subvillosum
C. tuvinicum 
 Thyrocarpus T. sampsonii
and Brachybotrys
(B. paridiformisChionocharis (C. hookeri and Trigono-
caryum (T. involucratum    
2016 ).






 







DNA sequencing and phylogenetic analyses. —





land 


Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
527Version of Record
TAX ON
65 (3) • June 2016: 523–546
 
  


trnL
and trnL-trnF  trnL-F
rps16trnS-G 




 



 

= 
 





 




 








Wellstedia Codon 

RESU LT S



 trnS-G trnL-F Andersonglossum occi-
dentale  and Decalepidanthus elongatus
 

  
trnL-Frps16
trnS-G
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
Decalepidan-
thus trollii 


  











  





 



Echiochi-
lon Antiphytum  





Caccinia
Trichodesma
  LasiocaryumChionocharis
hookeri and LasiocaryumMicrocaryum



Mertensia


Omphalodes



Asperugo pro-
cumbensMemoremea scorpioides
Mertensia
AnoplocaryumMerten-
sia 
  
Craniospermum 
 



Brachybotrys

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
528 Version of Record
TAX ON
65 (3) • June 2016: 523–546
Codon royenii W2373
Codon schenckii W2375 Wellstedia somalensis W3004
Wellstedia socotrana W3003
Memoremea scorpioides W2620
Heterocaryum subsessile W2378*
Suchtelenia calycina W3077*
Brachybotrys paridiformis W4053
Afrotysonia glochidiata W4239
Cynoglossum amplifolium W2980
Andersonglossum occidentale W4279*
Andersonglossum occidentale W0602*
Adelinia grande W2834*
Andersonglossum virginianum W0735*
Chionocharis
Anoplocaryum
Craniospermum
Trigonocaryum
Thyrocarpus
Brachybotrys
Afrotysonia
Craniospermeae
(Trichodesmeae p.p.)
Lasiocaryeae
(Mertensia-clade)
Asperugeae
(Omphalodes-clade)
Omphalodeae
(Eritri chieae s.str. plus *)
Rochelieae
(Core-Cynoglosseae)
Cynoglosseae
(Microula-clade)
Microulinae
(Cryptantha-clade plus *)
Amsinckiinae
(Bothriospermum-clade)
Bothriosperminae
(Cynoglossum s.l.)
Cynoglossinae
(Cynoglosseae s.l.)
Cynoglossoideae
(Echiochileae)
Echiochiloideae
Boraginoideae
Trichodesmeae
(Trichodesmeae p.p.)
Myosotideae
(Myosotideae)
Boragineae
(Boragineae)
Lithospermeae
(Lithospermeae)
Eritrichiinae
Heterocaryinae
Moritziinae
Boragininae
ML-BS
PP
= 100%
= 80%–99%
= 60%–79%
= 1.00

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
Version of Record
TAX ON
65 (3) • June 2016: 523–546
Tr igon ot is 

Decalepi-
danthus Myosotis 
Trigonocaryum
involucratum 


   
  Afro-
tysonia glochidiata  Cynoglossum amplifolium  
 
Microula  Metaeritrichium
microuloides Actinocarya tibetica 
Microula

Andersonglossum occidentale 

Andersonglossum virginianum 
Adelinia grandeHarpagonella
palmeri     Pectocarya Dasynotus
daubenmirei Cryptantha 
Plagiobothrys Amsinckia
AdeliniaAmsinckiaAndersonglossumCryptantha
DasynotusHarpagonellaPectocaryaPlagiobothrys
Cryptantha


Bothriospermum
Bothriospermum Thyro-
carpus sampsoniiNihon japonicum 
and N. akiensis  
Fig. .
Wellstedia and
Codon 




Fig. .



. .


86/1
Boraginoideae
Onosma rhodopaea W0582
Brunnera macrophylla W0628
Anchusa formosa W0781
Mairetis microsperma W0512
Echium creticum W2074
Moltkia petraea W0514
Pentaglottis sempervirens W0668
Lithospermum macbridei W1032
Onosma barsczewskii W4070
Symphytum cordatum W2749
Lithospermum peruvianum W1070
Arnebia linearifolia W2340
Echium vulgare W2063
Glandora nitida W1706
Lithospermum tschimganicum W1088
Moritzia lindenii W1977
Cystostemon hispidus W2719
Echium callithyrsum W2076
Lycopsis arvensis W0624
Lithospermum officinale W0409
Onosma tricerosperma W2125
Buglossoides incrassata W2304
Symphytum grandiflorum W2750
Cystostemon ethiopicus W2731
Pulmonaria angustifolia W2086
Trachystemon orientale W0666
Borago officinalis W2070
Antiphytum floribundum W2759
Antiphytum hintoniorum W2758
Echiochilon fruticosum W2542
Cerinthe major W1702
Wellstedia somalensis W3004
Wellstedia socotrana W3003
Codon schenckii W2375
Codon royenii W2373
0.98
79/1
87/1
90/1
95/1
97/1
97/1
0.97
74/1
(Cynoglosseae s.l.)
Cynoglossoideae
1
Echiochiloideae
(Echiochileae)
Boragineae
(Boragineae)
Lithospermeae
(Lithospermeae)
Boragininae
Moritziinae

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
530 Version of Record
TAX ON
65 (3) • June 2016: 523–546

Cynoglossum 
Cynoglossum LindelofiaMattiastrum
Microparacaryum Para-
caryumParacynoglossumPardoglossum 
RinderaSolenanthusTrachelanthus 
Cynoglossum 
Amsinckia Dasyno-
tusNihonPectocaryaPlagiobothrys 

Plagiobothrys

Fig. .       




 
Fig. .






(Cynoglosseae s.l. )
Cynoglossoideae
(continues in Fig. 5)
(Mertensia-clade)
Asperugeae
Omphalodeae
(Omphalodes-clade)
(Eritrichieae s.str. plus *)
Rochelieae
Myosotidium hortensia W0659
Caccinia macranthera var. glauca W4269
Hackelia velutina W1921
Hackelia bella W1110
Eritrichium canum W0343
Mertensia maritima W1230
Hackelia cf. andicola W2348
Hackelia sharsmithii W1111
Caccinia macranthera W2366
Mertensia ciliata W2728
Lasiocaryum munroi W2341
Hackelia thymifolia W2386
Microcaryum pygmaeum W2970
Austrocynoglossum latifolium W3079
Suchtelenia calycina W3077*
Mertensia lanceolata W2905
Mertensia sibirica W2622
Eritrichium pectinatociliatum W2978
Lappula squarrosa W2059
Hackelia deflexa W0480
Rochelia persica W2556
Eritrichium nanum W0588
Cynoglossum suaveolens W1491
Rochelia cancellata W1301
Eritrichium pamiricum W2343
Omphalodes verna W0627
Lappula shanhsiensis W2131
Trichodesma calcaratum W2362
Lappula sessiliflora W1279
Rochelia cardiosepala W1302
Trichodesma africanum W2363
Lappula occidentalis W1109
Eritrichium aretioides W2693
Asperugo procumbens W0004
Amblynotus rupestris W2294
Lappula marginata W2541
Memoremea scorpioides W2620
Heterocaryum subsessile W2378*
Caccinia strigosa W2055
Lasiocaryum trichocarpum W2603
Trichodesma zeylanicum W0163
Mertensia dahurica W2695
Omphalodes linifolia W0897
Omphalodes cappadocica W1893
Omphalodes aliena W2972
Omphalodes caucasica W2334
Omphalodes kuzinskyanae W2913
Myosotidium hortensia W2623
Omphalodes commutata W2128
Cynoglossum pauciflorum W3040
92/1
96/1
97/1
Chionocharis hookeri
K35577
89/0.96
Fig. 3
91/1
98/1 Anoplocaryum helenae W4037
Anoplocaryum compressum W4036
97/1
99/1
98/1
88/1
95/1
99/1
99/1 96/1
81
90/1
98/1
88/1
83/1
97/1
93/1
94/1
99/1
Fig. 5 (Cynoglossoideae, cont.)
74
75/0.98
94/1
Trichodesmeae
(Trichodesmeae p.p.)
Lasiocaryeae
(Tricodesmeae p.p.)
Eritrichiinae
Heterocaryinae

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
531Version of Record
TAX ON
65 (3) • June 2016: 523–546
Actinocarya tibetica W1139
Cryptantha granulosa W1912
Cryptantha pterocarya W2158
Decalepidanthus primuloides W2702
Cryptantha glomerata W2164
Lindelofia macrostyla W2336
Trigonotis jinfoshanica W2611
Decalepidanthus elongatus W2709
Trigonotis radicans W2354
Microula stenophylla W4282
Cryptantha paradoxa W2595
Trigonotis formosana var. elevatovenosa W1895 & W4273
Myosotis stolonifera W1229
Cryptantha johnstonii W2593
Pectocarya penicillata W2188
Dasynotus daubenmirei W2102
Pectocarya setosa W1919
Andersonglossum occidentale W0602*
Cryptantha texana W2757
Nihon akiensis W3038
Andersonglossum occidentale W4279*
Bothriospermum tenellum W4307
Trigonotis guilielmi W1124
Amsinckia vernicosa W2596
Metaeritrichium microuloides W2979
Myosotis incrassata W0378
Plagiobothrys congestus W2177
Andersonglossum virginianum W0735*
Cryptantha micrantha W2690
Harpagonella palmeri W2691
Myosotis caespitosa W1902
Paracynoglossum asperrimum W2115
Plagiobothrys linifolius W2180
Bothriospermum tenellum W2614
Plagiobothrys kunthii W2181
Amsinckia menziesii W2594
Pectocarya lateriflora W1920
Myosotis stenophylla W2114
Amsinckia tessellata W0276
Plagiobothrys humilis W2178
Dasynotus daubenmirei W2833
Microula filicaulis W4077
Cynoglossum amplifolium W2980
Cryptantha patagonica W1061
Decalepidanthus trollii 2703
Microula pustulosa W1129 & W4277
Amsinckia calycina W1918
Myosotis sparsiflora W0002
Nihom japonicum W3039
Plagiobothrys myosotoides W1685
Lindelofia olgae W2347
Cynoglossum magellense W1516
Rindera tetraspis W0753
Paracaryum rugulosum W1320
Cynoglossum officinale W0646
Cynoglossum montanum subsp. extraeuropaeum W1321
Paracaryum ancyritanum W1277
Paracaryum racemosum W2104
Cynoglossum germanicum W0487
Solenanthus apenninus W0654
Rindera schlumbergeri W2310
Cynoglossum clandestinum W1695
Cynoglossum creticum W0286
Mattiastrum lithospermifolium W2720
Pardoglossum cheirifolium W1709
Lindelofia longiflora W0663
Cynoglossum columnae W0285
Cynoglossum sphacioticum W1253
Cynoglossum dioscoridis W1856
Solenanthus tubiflorus W1693
Cynoglossum montanum W0321
Trachelanthus cerinthoides W2335
Paracaryum cappadocicum W1250
Cynoglossum troodi W1051
Microparacaryum intermedium W2981
Cynoglossum lanceolatum W3032
Cynoglossum coeruleum W4308
Cynoglossum amabile W4271
Cynoglossum australe W4272
Cynoglossum amabile W4270
(Core-Cynoglosseae)
Bothriospermum chinense W4050 & W4283
Craniospermeae
Fig. 4
97/1
Craniospermum tuvinicum W4055
Craniospermum sp. nov. W4041
Craniospermum subvillosum W4039
Craniospermum mongolicum W4042
Craniospermum echioides W4043
Brachybotrys paridiformis W4053
94/1
95/1
83/1
Trigonocaryum involucratum W4268
Afrotysonia glochidiata W4239
96/1
90/1 88/1
91/1
83/1
94/1
0.99
91/1
80/1
74/1
97/1
96/1
85/1 Thyrocarpus sampsonii W2598
95/1
0.95
1
0.99
1
0.97
99/1
0.95
0.98
89/1
99/1
Adelinia grande W2834*
(cont.) Cynoglossoideae
(Cynoglosseae s.l.)
(Cynoglossum s.l.)
Cynoglossinae
Bothriosperminae
(Cryptantha-clade plus *)
Amsinckiinae
70/1
100
96/1
97/1
0.95
Cynoglosseae
Microulinae
(Microula-clade)
Myosotideae
(Myosotideae)

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
532 Version of Record
TAX ON
65 (3) • June 2016: 523–546
DISCUSSION
Major clades of Boraginaceae. —




   


 


rps16trnL-F
trnS-G


Lasiocaryum  

Chionocharis 
  Microcaryum Lasiocaryum (Lasiocaryum



  





Lasiocaryum




 Anoplocaryum  
Mertensia

 Mertensia and Anoplocaryum
 Mertensia- 





Anoplocaryum
 Mertensia Asperugo
Asperugo Mertensia
Anoplocaryum Memoremea scorpioides 
Asperugo procumbens   

  
 

Memore-
mea scorpioides 
Omphalodes 
O. scorpioides 
 
 

O. scorpioides  

Memoremea

Omphalodes

Cranio-
spermum
   







Craniospermum and Trichodesma
 



 




   
 Brachybotrys and Trigonocaryum
Trigo notis
Myosotis 
Trigonotis and Brachybotrys   
Mertensia.
    

Brachybotrys
  

Tr igonot i s and
Myosotis 
. 
Trigonocaryum involucratum
Myosotis involucrata 
  
 
BrachybotrysTr igono t i s Myosotis
 Decalepidanthus (= Pseudomertensia; 
Myosotis
 



  


Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
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Afrotysonia
glochidiataCynoglossum amplifolium

  
MicroulaMetaeritrichium
  and Actinocarya 
Microula  
 
Afrotysonia and Microula
 
Fig. . Craniospermum. A–D, Craniospermum canescens 
W. Hilbig, D. Bumschaa & al. 104/79   E & F, C. mongolicumE. Jäger F225 A, B, C, E & F,
D,

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
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65 (3) • June 2016: 523–546

 

Microula
 

Afrotysonia 

Afrotysonia 
   
   
Afrotysonia   
 
 Afrotysonia-Microula
CryptanthaAnder-
songlossum and Adelinia
Andersonglossum  
 
Cryptantha
AmsinckiaCryptantha
Cryptantha
Dasynotus HarpagonellaPecto-
caryaPlagiobothrysAmsinckia and
Plagiobothrys  Cryptantha
Cryptantha  

   
Cryptantha
  Cryptantha
Harpagonella

Cryptantha
 
Plagiobothrys
     

 Thyro-
carpus  Nihon
Bothriospermum chinense B. tenellum
Bothriospermum
Bothriospermum
and Thyrocarpus Thyrocarpus
Bothriospermum 

 
Bothriospermum B. secundum
B. chinense and B. tenellum 
  Thyrocarpus sampsonii. Thyrocarpus 
Antiotrema 
Bothriospermum

  
 Nihon akiensis and N. japonicum 
in Omphalodes) 
Bothriospermum 
Bothriospermum and Thyrocarpus
 Nihon 
Antiotrema Bothriospermum and
Thyrocarpus 
  
.
CynoglossumCyno-
glossum  

 
Lindelofia Mattiastrum
Microparacaryum Paracaryum Pardoglossum Rindera So-
lenanthus and Trachelanthus 
 Cynoglossum 

Main subdivi sions in Cynoglosseae s.l. and t he placement
of the problematic genera. —


 



   




Omphalodes


     Caccinia
Trichodesma   Lasiocaryum Chionocharis
LasiocaryumMicrocaryum


 
Lasiocaryum
 AsperugoMemo-
remea and Mertensia. Anoplocaryum  
Mertensia
  Mertensia 

      Hetero-
caryumSuchtelenia 
Eritrichium HackeliaLappula
Rochelia

    


Brachybotrys Tr igo
-
notis and Decalepidanthus Myosotis
Trigonocaryum  Myosotis.  

Trigonotis radicans 

Brachybotrys 
Omphalotrigonotis 
 

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
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65 (3) • June 2016: 523–546

Mertensia Omphalodes 

Bothriospermum
 Ogastemma
Sericostoma
Moltkiopsis Neatostema




Afrotysonia glochidiata and Cynoglossum amplifolium
Microula Actinocarya
and Metaeritrichium 
Cynoglossum
and Afrotysonia 
 Cryptantha
 
Cryptantha 

Cynoglos-
sum   
 
Cynoglossum Cryptantha


Cryptan-
tha 
Cryptantha


      Bothriospermum
Cynoglossum

Bothriospermum Thyrocarpus samp-
sonii Nihon akiensis and N. japonica.

SinojohnstoniaAntiotrema
 Antiotrema 
Bothriospermum and Thyrocarpus).



FORMAL TAXONOMY










 

    
 

Conspectus of the infrafamilial classification of
Boraginaceae
 
 
  
   
   
  
 
  
  
  
  
  
   
   
  
  
  
   
   
   
   
Echiochiloideae  subfam . no v.
 
       

Echiochilon

Antiphytum 
 AntiphytumAmblyno-
topsisAmphibologyne Chamis-
soniophila EchiochilonChil-
echium Chilochium     Echiochilopsis
ExioxylonLeurocline
SericostomaTetraedrocarpus  
Ogastemma    Megastoma  

Megastoma

    
    
 

  
 
  
 


Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
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65 (3) • June 2016: 523–546





 
 

Antiphytum and Ogastemma
Echiochilon
Antiphytum



 
 



Boraginoideae   
Bora geaeBorago
 
       
  
          


Tab le . 
   


 Antiphytum  Echiochilon  Ogastemma 30
   Anchusa  Anchusella Borago  Brunnera  Cynoglot-
tis Gastrocotyle Hormuzakia Lycopsis  Melanortocarya Nonea
Pentaglottis  Phyllocara Pulmonaria  Symphytum
 Trachystemon 
140
 Moritzia  Thaumatocaryon 6
 Aegonychon Alkanna Ancistrocarya Arnebia  Buglossoides
 Cerinthe  Cystostemon  Echiostachys Echium
 Glandora  Halacsya Huynhia Lithodora Lithosper-
mum  Lobostemon Maharanga Mairetis  Moltkia 
Moltkiopsis Neatostema Onosma  Paramoltkia Podonosma
Pontechium Stenosolenium
460
  Caccinia  Trichodesma  50
 Chionocharis  Lasiocaryum  Microcaryum  5
 Anoplocaryum  Asperugo  Memoremea  Mertensia

50
 GyrocaryumIberodes Mimophytum Myosotidium Omphalodes
 SelkirkiaMapuchea
35
  Eritrichium  Hackelia  Lappula  Lepechini-
ella Rochelia 
200
 Heterocaryum  Suchtelenia  7
 Craniospermum  6
 Brachybotrys  Decalepidanthus  Myosotis 
Trigonotis  Omphalotrigonotis
160
  Cynoglossum  Microparacaryum Lindelofia
 Mattiastrum  Paracaryum  Rindera
 Solenanthus
200
 Antiotrema Bothriospermum  Nihon  Thyrocarpus 
Sinojohnstonia
14
 Afrotysonia  Microula Adelocaryum36
 Amsinckia  Cryptantha  Eremocarya Greeneo-
charis Johnstonella Oreocarya Dasynotus  Harpagonella
 Nesocaryum Pectocarya  Oncaglossum Plagiobothrys
 Adelinia  Andersonglossum 
300




Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
537Version of Record
TAX ON
65 (3) • June 2016: 523–546

 


Boragineae
Borago
 
Anchusa
  
Symphytum




 


  






 Thaumato
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caryon and Moritzia







Boragininae
Borago
  Anchuseae
Anchusa
 Pulmonarieae
Pulmonaria

SymphyteaeSymphytum
 Anchusa Buglossum
Anchusella Borago
BuglossitesBrunneraCynoglottis
GastrocotyleHor-
muzakiaLycopsisBuglossa
EchioidesOskampia
OskampiaMelanortocarya
 Nonea Elizaldia
MassartinaNephrocarya
   Pentaglottis Caryolopha
Phyllocara
PulmonariaBessera Paraskevia
SymphytumProcopi-
ania ProcopiphytumTrachystemon
NordmanniaPsilo-
stemon




 


   



 


 




 

Moritziinae  subtr. nov. Moritzia

MoritziaMeratia
MeratiaThaumatocaryon


  




   



     
  

 
M. lindenii 




Lithospermeae
Lithospermum

Alkanna
        
Cerinthe

EchieaeEchium

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
538 Version of Record
TAX ON
65 (3) • June 2016: 523–546

Moltkiopsis 
AegonychonMargarospermum
RhytispermumAlkanna 
1824 (incl. BaphorhizaCamptocarpus
CampylocaryumOnochilis
Ancistrocarya    Arnebia   
Arnebiola   Dioclea   Leptanthe
    Macrotomia   Meneghinia 
MunbyaStrobilaTeta r i s
Tox o stigmaBuglossoides
Cerinthe CystostemonVaupelia
Echiostachys Echium
(incl. ArgyrexiasIsoplesionLarephes
MegacaryonStomotechium
 Glandora     
Halacsya Zwackhia  Huyn-
hia AipyanthusEchioides
    Lithodora Lithospermum 
1753 (incl. BatschiaCyphorima
LasiarrhenumMacromeria
NomosaOnosmodiumOsmo-
dium Pentalophus
PerittostemaPsilolaemus
PurshiaPurshia
Ulugbekia Lobostemon 
EchiopsisIsoriumLobostema
 OplexionPenthysaTraxara
MaharangaMairetis  Molt-
kiaGymnoleimaMoltkiopsis
NeatostemaOnosma
1762 (incl. ColsmanniaCoriantha
Choriantha Sava   Paramoltkia 
 PodonosmaPontechium 
Stenosolenium



 
Cerinthe 
  
  





  
 

      
   




       
   
Ancistrocarya MaharangaEchiostachys
LobostemonLithospermum


   
  Lithospermum and in Echium





Cynoglossoideae subf am. n ov.
Cynoglossum

 

 
 

 
 


    




 

 

 

 








  

  Trichodesmeae  
Trichodesma 
 CacciniaAnisanthera
HeliocaryaTrichodesma
(incl. Boraginella Boraginodes
BorraginoidesFriedrichsthalia
Lacaitaea Leiocarya 
OctosomatiumPollichia         
SpiroconusStreblanthera

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
Version of Record
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65 (3) • June 2016: 523–546






  





Trichodesma scottii

TrichodesmaSolanumCac-
cinia
Lasiocaryeae  t r. nov. Lasiocaryum

 Chionocharis Lasiocaryum
   Oreogenia    
OrogeniaSetulocarya R.R.Mill &
 Microcaryum
   
       
        
    








Asperugeae 
 
Asperugo
    
Anoplocaryum
AnoplocaryumAsperugo
Memoremea 
Mertensia Casselia 
Cerinthodes HippoglossumOreo-
charisPlatynemaPneumaria
SteenhammeraWinkleria
        






   



Mertensia
   
Asperugo
and MemoremeaMertensia
Anoplocaryum
Memoremea have


Omphalodeae tr. nov.Omphalodes
Mill.
GyrocaryumIberodes 
     Mimophytum   
Omphalodes Mill. 1754 (incl. Omphalium 
Picotia  Umbilicaria
Myosotidium   Selkirkia
Mapuchea 
2016 ).
 













Rochelieae Rochelia

    
Eritrichium
=      
 Hetero-
caryum

   
        

   
 





 



 Eritrichium

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
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65 (3) • June 2016: 523–546
Hackelia 


Eritrichiinae
 Eritrichieae   Eritrichium


  Echinospermum  

  
Rochelia
 Eritrichium Ambly-
notusSauriaTianschaniella
Hackelia Austrocynoglos-
sumEmbadiumLap-
pula  Cynoglossospermum
EchinospermumOmphalolappula
SclerocaryopsisLepechiniella 
Rochelia Cervia
Raclathris




  
 






Rochelia


  Eritrichium
  Hackelia



Heterocaryinae
Heterocaryum
HeterocaryumSuchtelenia

 
 
 
  



 





  

Craniospermeae

(‘Craniospermeae’Craniospermum
 Craniospermum Diploloma
 



 



Craniosper-
mum


Myosotideae

MyosotidesMyosotis

Pseudomertensia Riedl.

Trigonocaryum
 
Zoelleria
 Brachybotrys     Deca-
lepidanthus Pseudomertensia
Scapicephalus   Myosotis  
(incl. Echioides  Echioides
  ExarrhenaGymnomyosotis
ScorpioidesStrophi-
ostomaTrigonocaryum Trig onot i s
 EndogoniaHavilandia
PedinogyneStephanocaryum
Zoelleria*Omphalotrigonotis    

 
   


   






Myosotis  
  Omphalotrigo-
notis    
Trigonotis and Myosotis

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
541Version of Record
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65 (3) • June 2016: 523–546


Cynoglosseae 
Cynoglossum
= 
Harpagonella
  
Cryptantha


 








CynoglossinaeCyno-
glosseae Cynoglossum

Rindera
Cynoglossum 
AnchusopsisBilegnumCerinthopsis
CrucicaryumCynoglos-
sopsis   CyphomattiaIvanjohnstonia
Kuschakewiczia*Linde-
lofia Mattia*Mattiastrum 
  *Paracaryum   Paracynoglossum
 Pardoglossum*Rindera
*Solenanthus  *Trachelanthus 
 Microparacaryum
Brandella
 






 





   
Microparacaryum
Cynoglossum 
          
Cynoglossum
   
  
in Cynoglossum.

  



 Bothriosperminae 
 Bothriospermum 
Antiotrema Henryettana
Bothriospermum Nihon 
 Thyrocarpus 
Sinojohnstonia














 

Sinojohnstonia
 Microulinae s ubt r. nov.Micro-
ula 
Afrotysonia Tysonia
TysoniaMicroula
1876 (incl. Actinocarya Glochidocaryum  
MetaeritrichiumSchistocaryum
Tretocarya*Adelocaryum 
(incl. Paracaryopsis












Afrotysonia


AfrotysoniaCynoglossum
Adelocaryum
Cynoglossum amplifolium


Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
542 Version of Record
TAX ON
65 (3) • June 2016: 523–546
3.8.4 S Amsinckiinae
AmsinckieaeAmsinckia


Harpagonella

Allocarya
 *Adelinia      *Andersonglossum
Amsinckia Cryptan-
tha Eremocarya  
GreeneocharisHemisphaerocarya
 JohnstonellaKrynitzkia
     Oreocarya Piptocalyx
WheelerellaDasynotus 
Harpagonella Nesocaryum
Pectocarya Ctenospermum 
GruveliaKtenospermum
Oncaglossum      Plagiobothrys 
Allocarya  Allocaryastrum
 Echidiocarya
EchinoglochinGlyptocaryopsisMac-
coyaSonnea
  




  

  
    
         

    

   Plagiobothrys


 


Pectocarya

Amsinckia Cryptantha Plagiobothrys 



  

Dasynotus dauben-
mirei and Cynoglossum 
  
Andersonglossum
OncaglossumDasynotus un
Cryptantha 
  





ACKNOWLEDGEMENTS
       

Afrotysonia pilosicaulis
Craniospermum 
   
Afrotysonia glochidiata 




 Lasiocaryum and Chionocha-
ris
Craniospermum
  Cynoglossum
 

 

 

 

LITERATURE CITED
Avetisjan, E.M.   
Trudy Bot. Inst. Akad. Nauk Armen. S.S.R. 10: 1– 66.
APG III 
 
Bot. J. Linn. Soc. 161: 10 5 –121.
           
Bentham, G. 
Genera plantarum(2) . : 
Reeve.

Bolus, H. Tysonia africanaHooker’s Icon.Pl. 20.

Brand, A. :  
   Das Pflanzenreich   252. 

Cecchi, L . & Selvi, F. 
Halacsya and Paramoltkia   

matKTaxon 58: 700–714 .
Cohen, J.I. ecu
 
   Cladistics 
      
Cohen, J.I. 2015. Adelinia and Andersonglossum 
Cynoglossum. Syst. Bot.
 
Dickoré, W.B. & Hilger, H.H. 2015.
Decalepidanthus


Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
543Version of Record
TAX ON
65 (3) • June 2016: 523–546
C.M., Hasenstab-Lehman, K.E., Hilger, H.H., Miller, J.S.,
Mittelbach, M., Nazaire, M., Nepi, M., Nocentini, D., Ober, D.,
Olmstead, R.G., Selvi, F., Simpson, M.G., Sutorý, K., Valdés,
B., Walden, G.K. & Weigend, M.  
Taxon 65: 502–522.
 
Luo, D., Yue, J.-P., Sun, W.-G., Xu, B., Li, Z.-M., Comes, H.P. &
Sun, H.   

J. Biogeogr. 43: 31– 43.
      
Mansion, G., Selvi, F., Guggisberg, A. & Conti, E. 
  
       
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         
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Mertensia. Syst.
Bot. 37: 758–783.      
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Lithodora
NeatostemaMairetisHalacsya
and ParamoltkiatrnLUAA
Tax o n
Van Royen, P. 
aPacific Sci.
Verdcourt, B.  . 1–124 in:  
Flora of tropical East Africa
Weigend, M., Got tschling, M., Selvi, F. & Hilger, H.H. 
Lithospermum
          
Molec. Phylogen. Evol. 52: 755–768.
 
Weigend, M., Gottschling, M., Selvi, F. & Hilger, H.H.  
  
 Syst.
Bot.  
Weigend, M., Luebert, F., Selvi, F., Brokamp, G. & Hilger, H.H.
Cynoglossum
Omphalodes Mill.) –  
Molec. Phylogen. Evol. 68: 604–618 .
 
Weigend, M., Luebert, F., Gottschling, M., Couvreur, T.L.P., Hilger,
H.H. & Mi ller, J.S.     –
Cladistics 30: 50 8 –518.
      
Weigend, M., Selvi, F., Thomas, D.C. & Hilger, H.H. 2 016 . 
  –102 i n:     The families
and genera of vascular plantsFlowering plants: Eudicots;
Aquifoliales, Boraginales, Bruniales, Dipsacales, Escalloniales,
Garryales, Paracryphiales, Solanales (except Convolvulaceae),
Icacinaceae, Metteniusaceae, Vahliaceae. 
. 
Wiens, J. J. 
Syst. Biol. 47: 568–581.
 
Zhu, G.-L., Riedl, H. & Kamelin, R.     
Flora of China16

Appendix 1.
trnL-Fr ps16trnS-G

Actinocarya tibetica    Sino- British Qinghai Ex pedition 1997-658Adelinia grande  
M. SchwarzländerAfrotysonia glochidiata
Bräuchler & Carbutt 6423  Amblynotus rupestrisHilg er 159 7
Amsinckia calycina  Weigend & Schwarzer 8031 
Amsinckia menziesiiWeigend 9010Am-
sinckia tessellata Hoffmann 4/98Amsinckia vernicosa Weigend
9008Anchusa formosa Bigazzi & Selvi 97.006
Andersonglossum occidentaleNelson s.n.
Andersonglossum occidentaleDavis & Lightowlers 66961Anderson-
glossum virginianumNe l son 21124 Anoplocaryum compressum     
Chan & Balde s.n.Anoplocaryum helenaeGolajkov s.n.
Antiphytum floribundumLott & al. 5574
Antiphytum hintoniorum Patterson & al. 1415 
Arnebia linearifolia Shahin Zarre & al. 528 Asperugo procumbens Cult. Berlin,
Hilger s.n. Austrocynoglossum latifoliumCult. Berlin, Weigend 9441
Borago officinalis Erixon & Bremer 11
Bothriospermum chinenseA. Yu. Korolyuk, E.A. Korolyuk s.n. Both-
riospermum tenellumPeng Li ZC-2  Bothriospermum tenellum
Cult. Bot. Ga rd. Bonn (ID 10691)Brachybotrys paridiformis 
Ovchinnikova s.n.Brunnera macrophyllaHilger
s.n.Buglossoides incrassataSelvi & al. 07.40 
Caccinia macranthera   Shahin Zarre 700 Caccinia
macranthera glauca    Lobin & Weigend 221-1 Caccinia strigosa 
Mohr M592Cerinthe majorBigazzi & Selvi 04.22
Chionocharis hookeri  Cra wford & al. 571    Codon royenii
Greuter 21551     Codon schenckiiWalter & Walter 118 
Craniospermum echioides   A.Yu. Korolyuk s.n. 
Craniospermum mongolicumA.Yu . Korolyu k s.n.Cranio-
spermum sp. nov.,A.Yu. Korolyuk s.n.Craniospermum subvillosum
Kovto nyu k 741 Craniospermum tuvinicumChan & Balde s.n.
Cryptantha glomerataWeigend & Hilger 5935
Cryptantha granulosaWeigend & al. 2000/642 
Cryptantha johnstoniiCult. Berlin, Weigend 9007 Cryptantha micrantha
Cult. Berlin, Weigend 9058 Cryptantha paradoxa 
Cult. Berlin, Weigend 9136Cryptantha patagonicaWeigend & al.

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
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65 (3) • June 2016: 523–546
5957Cryptantha pterocaryaFah l H 20 a/199 8
Cryptantha texanaCa rr 25514Cyno-
glossum amabile Cult. Bot. Gard. Bonn, Joßberger P663  
Cynoglossum amabileCult. Bot. Gard. Bonn, Joßberger, P402  
Cynoglossum amplifoliumCult. Bot. Gard. Bonn, Koelker 06Cynoglos-
sum australe   Cult. Bot. Gard. Bonn, Joßberger P648 Cynoglossum clandestinum
Bigazzi & Selvi 04.21Cynoglossum coeruleumCult. Bot. Gard. Bonn
(ID 34835)Cynoglossum columnae Hilger 98/7 
 Cynoglossum creticumHilger 98/9 Cynoglossum dioscoridis
Biga zzi & Selvi 05.33 Cynoglossum germanicumHilger s.n.  
Cynoglossum lanceolatumCult. Berlin, Weigend 9198
Cynoglossum magellenseBigazzi & Selvi 03.05Cynoglossum montanum
H ilg er 97/17 Cynoglossum montanumextraeuropaeumBigazzi &
Selvi 02.23Cynoglossum officinaleHilger s.n.
Cynoglossum pauciflorum Kern 14 .12. 2009 Cynoglossum
sphacioticumHilger s.n.Cynoglossum suaveolens   Lepschi
& Craven 3937Cynoglossum troodiBrullo & al. s.n.
Cystostemon ethiopicus Thulin & al. 3576  
 Cystostemon hispidusForstreuter 91702
Dasynotus daubenmireiCrockett 492Dasynotus daubenmirei
Schwarzländer s.n.Decalepidanthus elongatus
Nü sser 874Decalepidanthus primuloidesM. Ackermann s.n. ED19751894
Decalepidanthus trolliiCult. Edinburgh, Sherriff s.n. ED19391024
 Echiochilon fruticosumKagiampaki s.n. 
Echium callithyrsumHilger 92/3482Echium creticumFörther s.n.
Echium vulgare  Moh r 597Eritrichium
aretioidesWeigend 9126Eritrichium canumFörther
s.n.  Eritrichium nanumZippel & al. s.n.  
Eritrichium pamiricum Anders 8098Eritrichium
pectinatociliatumBoufford & al. 29997Glandora nitida   
Thomas 1-05Hackelia bellaMerello & al. 702
Hackelia cf. andicolaWeigend & Bro kamp 8847   
Hackelia def lexaFrey s.n.Hackelia sharsmithii
Hilger U.S.A. 94/18   Ha ckelia thymifoliaHurk a 10234 
Hacke lia velutinaHilger 411/1997
Harpagonella palmeriWeigend 9056 Heterocaryum subsessile
Rechinger 33789 Lappula marginata Hilg er 1616 
Lappula occidentalisHilger 96/2
Lappula sessiliflora    Bigazzi & Selvi 02.35Lappula shanhsiensis    
Kürschner & al. 634 Lappula squarrosa M oh r 591
Lasiocaryum munroiPoelt 276
Lasiocaryum trichocarpumSinclair & Long 5405 Lindelofia
longifoliaWeigend 9120 Lindelofia macrostyla
Vas ak s.n.  Lindelofia olgae Kaletkina s.n. 17.06.1966
Lithospermum macbrideiWeigend & al. 5073
Lithospermum officinaleWerres & Ristow s.n.  Lithospermum peruvianum
Weigend & al. 2000/761Lithospermum tschimganicumOrazowa & Fissjun 5785
Lycopsis arvensisHilger & Werres s.n. 
Mairetis microspermaPodlech 48277Mattiastrum lithospermifolium
Bergmeier & Matthäs 3422  Memoremea scorpioides   
Weigend 9039 Mertensia ciliataAckermann
709Mertensia dahuricaHil ge r 1601
Mertensia lanceolata Weigend 9171Mertensia maritima
Hilger s.n.Mertensia sibiricaWeigend 9069   
Metaeritrichium microuloides Boufford & al. 32025 Microcaryum
pygmaeumBoufford & al. 36208 Microparacaryum intermedium
Kü rschn er 10 -52        Microula filicaulis 
Boufford 40060     Microula pustulosaSino-American-British
Yushu Expedition (1996) 2328 Mic roula stenophylla  Zang & al. 121
Moltkia petraeaHilger s.n. 
Moritzia lindeniiE rik se n 59018 Myosotidium hortensia
Hilger s.n.Myosotidium hortensiaWeigend 9068 
Myosotis caespitosaHilg er 1575    
Myosotis incrassataHilger 98/5Myosotis sparsiflora
Weigend 8138Myosotis s tenophyllaM ohr 614
Myosotis stoloniferaHilger s.n.Nihon
akiense         Suga s.n.Nihon japonicum 
Kubota s.n.Omphalodes aliena
Hinton 28565Omphalodes cappadocicaCult. Berlin,
Weigend 8196Omphalodes caucasicaCult. Munich, Anon ann o 1969
Omphalodes commutataHilger s.n.Ompha-
lodes kuzinskyanaeM ohr 611 Omphalodes linifoliaHilger
Appendix 1. 

Chacón & al. • Taxonomic subdivision of Boraginaceae s.str.
546 Version of Record
TAX ON
65 (3) • June 2016: 523–546
s.n.  Omphalodes verna Hilger s.n.
 Onosma barsczewskiiOvchinnikova 157 Onosma rhodopaea
Hil ger 97/ 24Onosma tricerosperma  Hilger s.n .
Paracaryum ancyritanum Bigazzi & Selvi 02.39Paracaryum
cappadocicum Bigazzi & Selvi 02.41Paracaryum racemosum 
Tarikahya 1537aParacaryum rugulosum  Bigazzi & Selvi 02.42
 Paracynoglossum asperrimum Mo hr 615 
  Pardoglossum cheirifolium Bigazzi & Selvi 04.25
Pectocarya lateriflora Weigend & Schwarzer 7810 Pectocarya peni-
cillata   Fa hl H19/1998  Pectocarya setosa Hilger
14/199 8Pentaglottis sempervirensSelvi & Bigazzi 6/2000
Plagiobothrys congestus Villavicencio s.n.
Plagiobothrys humilisWeigend & Schwarzer 8075
Plagiobothrys kunthii   Weigend & Schwar zer 7832   Plagiobothrys linifolius
Schwarzer 04Plagiobothrys myosotoides
Weigend & Schwarzer 7825Pulmonaria angustifolia M. & K. Weigend 1999/45
Rindera schlumbergeriSelvi & al. 07.32
Rindera tetraspis  Sagalaev & Rusa novich 18328Rochelia cancellata
 Bigazzi & Selvi 02.53Rochelia cardiosepala Bigazzi & Selvi
02.61Rochelia persica   Rechinger 10377  
Solenanthus apenninus Frey s.n.  Solenanthus tubiflorus
Bigazzi & Selvi 04.23Suchtelenia calycinaParolly &
al . 130 80Symphytum cordatumCult. Be rlin, Weigend 8047
  Symphytum grandif lorumCult. Berlin, Weigend 9034
Thyrocarpus sampsonii   Liu Zh eng-yu 15360Trachelanthus cerinthoides
Parishani 14275Trachystemon orientalisCult. Berlin,
Weigend 9030 Trichodesma africanum Podlech 35050  
  Trichodesma calcaratum Podle ch 41053  Trichodesma
zeylanicumSchultka 12Trigonocaryum involucratum  
Gröger & al. 213-20   Trigonotis formosana va r. elevatovenosa 
  Cult. Berlin, Weigend 8128  Trigonotis guilielmi Azu ma 2001 
 Trigonotis jinfoshanica Peng Li ZC-?-2
Trigonotis radicansStarczenko & Ivanova 5488Wellstedia so-
cotranaThulin & al. 10155Wellstedia somalensis
Thulin & al. 10084
Appendix 1. 