ArticlePDF Available

Generic phylogeny and character evolution in Urticeae (Urticaceae) inferred from nuclear and plastid DNA regions

Authors:
  • Honam National Institute of Biological Resouces

Abstract and Figures

This study examines phylogenetic relationships among the 12 genera of Urticeae (Urticaceae) and investigates the pattern of morphological evolution based on analysis of nuclear ribosomal internal transcribed spacer (nrITS) and two plastid DNA regions (rbcL exon, trnL-F spacer). Sequence data were analyzed using maximum parsimony and Bayesian inference, and selected morphological traits were mapped onto the molecular tree. The molecular results strongly supported monophyly of Urticeae, excluding Gyrotaenia, which is related to Elatostemateae. All genera were monophyletic except for Urtica, Laportea, and Urera. Two Hesperocnide species are nested within Urtica. Laportea and Urera are divided into three groups with a strong geographical signal. The inferred phylogeny indicates five well-supported clades in Urticeae: clade A including Urtica (with Hesperocnide), Zhengyia, Laportea I, and Nanocnide; clade B comprising Dendrocnide and Discocnide; clade C including only Girardinia; clade D including Laportea II; and clade E including Obetia, Urera I, II, III, and Poikilospermum. Although it is difficult to identify morphological synapomorphies for these well-defined clades within Urticeae, character analysis shows that the herbaceous habit and alternate leaves are the ancestral states in the tribe. The presence of stinging hairs is the derived status in Urticeae, and it might have been a key innovation triggering species diversification in the tribe.
Content may be subject to copyright.
65
Kim & al. • Phylogeny and character evolution of Urticeae
TAX O N
 
Version of Record

;  || 
INTRODUCTION
 

    
 
 
Lam.
& 




 
Poikilospermum 
         
        

Poikilospermum
        
   



   
 



   
Gyrotaenia
      
Generic phylogeny and character evolution in Urticeae (Urticaceae)
inferred from nuclear and plastid DNA regions
Changkyun Kim,1* Tao D e ng,1,2,3* Mark Chase,4 Dai-Gui Zhang,5 Ze-Long Nie1 & Hang Sun1
 Key Laboratory for Plant Diversity and Biogeography of East Asia, Kunming Institute of Botany, Chinese Academy of Sciences,

  
 
 
  

 


DOI
   
Abstract
        
   
rbcLtrnLF
 
GyrotaeniaLaportea
      . Laportea           

ZhengyiaLaportea Nanocnide Dendrocnide Discocnide 
GirardiniaLaportea ObetiaPoikilospermum
 


Keywords

Supplementary Material


66
TAX O N
 Kim & al. • Phylogeny and character evolution of Urticeae
Version of Record

Girardinia

 






     

       
rbcLtrnL    

Dendrocnide       
Discocnide  Laportea  
  


  
     
Laportea
ObetiaPoikilosper-
mum
LaporteaL. cuspidata
Zhengyia
   
Table 1.
 

Dendrocnide
.
Discocnide

Girardinia

Gyrotaenia



Laportea

      
      
      
      
      
      
 

    

 37   
 



mala



 






Nanocnide

Obetia

Poikilospermum


  L.
Zhengyia

   

  
      
      
    

 
      
      
 


 







  7 35  
  













67
Kim & al. • Phylogeny and character evolution of Urticeae
TAX O N
 
Version of Record
     


Nanocnide  Obetia
    


  
NanocnideGirardinia   
PoikilospermumDendrocnide
 




 
DendrocnideDiscocnideGirardiniaGyrotaeniaLaportea
NanocnideObetiaPoikilospermaZhengyia
 Zhengyia Nanocnide
 
  








rbcLtrnLF
    
      

     

MATERIALS AND METHODS
Taxon sampling. —
 



        
DendrocnideDiscocnideGirardinia
GyrotaeniaLaporteaNanocnideObetiaPoi-
kilospermum
      
Zhengyia
   
     

        
trnLF  
   Fatoua villosa 
Morus albaSorocea affinis 
   
Cannabis sabiva
lupulus  
rbcLtrnL-F

DNA isolation, PCR amplification, and sequencing. —



        
rbcL 
  
  
 rbcL  
 
   trnLF  

          
  

   




  
      

Phylogenetic analyses. —
   


     




trnL-F


   
 

 

     
     


     


   
  Laportea


   *  
   
    

TAX O N
 Kim & al. • Phylogeny and character evolution of Urticeae
Version of Record
Fig. 1.
 
A–E,

A,
Obetia radula 
B,
Poikilospermum
C,
Dendrocnide
amplissima 
D,
Zhengyia shennongensis
E,

F–G,
  
F,
Laportea cus-
pidata
G,
    
H,
Girardinia suborbiculatatriloba  
I–J,

I,
Zhengyia shen-
nongensis;
J,
Laportea bulbifera.
K–M,

K,
Poikilospermum
L,

 
M,

N,
Zhengyia shennongensis  
O–Q,

Dendrocnide basirotunda
P,
Laportea bulbifera
Q,
 
69
Kim & al. • Phylogeny and character evolution of Urticeae
TAX O N
 
Version of Record
    
   
 


       

  
    



     
      




Morphological character evolution. —


   


     


   
         
   


      






  


RESULT S
ITS and plastid analyses. —
 
    



         

 


 Gyrotaenia.
Gyrotaenia myriocarpa 


Laportea
. Laportea


rbcLtrnLF

 


Girardinia 
DiscocnideDendrocnide 
 ZhengyiaLaportea I–Nanocnide 
Gyrotaenia
Laportea
 
Laportea  
   
  Laportea
ZhengyiaLaportea I–Nanocnide

 Laportea II




Table 2.


  rbcL trnLF 
     
  937   
     
     
  96  95 
     
     
     
     

TAX O N
 Kim & al. • Phylogeny and character evolution of Urticeae
Version of Record
Combined data analysis. —
  
          

    
Gyrotaenia myriocarpa

 Laportea
 
   
  ZhengyiaLaportea I–
Nanocnide  DendrocnideDiscocnide
Girardinia
 Laportea Obetia
Poikilospermum
Zhengyia shennongensis     
 
  Laportea  Nanocnide
Obetia
   
Poikilospermum


Morphological character evolution. —
   
   


       
   Dendrocnide
Discocnide  ObetiaPoikilospermum
       



          



Poik ilospermum

 


    
    

DISCUSSION
       





Phylogenetic position of Gyrotaenia. —
 
  Gyrotaenia 

Gyrotaenia

  

Gyrotaenia   
 

Gyrotaenia


                        

         Poi-
kilospermumGyrotaenia

Gyrotaenia
Elatostema
     Gyrotaenia
MyriocarpaPileaLecanthus  
ElatostemaGyrotaenia
     
 
Gyrotaenia 
Polyphyletic genera within Urticeae. —

DendrocnideDiscocnideGirardinia
NanocnideObetiaPoikilospermumZhengyia

Laportea

     
     




                  

  

    
     
 



  

Laportea  
    

         
      

Laportea
     


Kim & al. • Phylogeny and character evolution of Urticeae
TAX O N
 
Version of Record
Fig. 2.




TAX O N
 Kim & al. • Phylogeny and character evolution of Urticeae
Version of Record
Fig. 3.

        
                   

73
Kim & al. • Phylogeny and character evolution of Urticeae
TAX O N
 
Version of Record
-
pidata    

Laportea     

LaporteaSceptro-
cnide Fleurya  
Laportea
    
   

Laportea Sceptrocnide 


Laportea    
Laportea 
   -
sisFleurya
Laportea                   
   


  Laportea
Laportea
Laportea 



Laportea    
     
         
 ZhengyiaLaportea I–Nanocnide
Laportea II–Obetia
Poikilospermum       


   
 
   
      
   
Laportea   
       
  

ObetiaPoikilospermum  

Laportea      
  
     





    




    
  
 
   
 
   




 

 
    



    

  

 




Phylogenetic relationships within Urticeae. —



  
Zhengyia  Nanocnide–Laportea cus-
pidata   Zhengyia 
 
 
    

Zhengyia   

    
 Zhengyia  
     
   


Zhengyia

al
Nanocnide  
 


 

 Nanocnide   

TAX O N
 Kim & al. • Phylogeny and character evolution of Urticeae
Version of Record

Laportea cuspidataLaportea
       Girardinia    
Laportea
   Girardinia  
   
 GirardiniaZhengyia   

 
  
  
Nanocnide
DendrocnideDiscocnide 

      
Dendrocnide


 Dendrocnide
   Dendrocnide 
Discocnide Dendrocnide

Discocnide
  
  

Girardinia diversifolia
    
     
Laportea
          
    
Girardina  Dendrocnide
Discocnide Laportea     
Girardina DendrocnideDiscocnide

  
   
diversifolia  
triloba     
     suborbidulata  

    

 diversifolia



       Laportea II
ObetiaPoikilospermum

Laportea   

    
    Obetia   
Poikilospermum   III. 
ObetiaLaportea
Girardinia
  Obetia 





Poikilospermum         
      
  
 


   
Poikilospermum
 
 
      

Poikilospermum
Poikilospermum



Poikilospermum   

 
  

Urticeae 
Poikilospermum.
 DendrocnideDiscocnide Girardinia
LaporteaNanocnideObetiaPoikilospermum
 Zhengyia.
Gyrotaenia.
Morphological character evolution. —
  


Zhengyia 


     



    


    
     


 


75
Kim & al. • Phylogeny and character evolution of Urticeae
TAX O N
 
Version of Record
Akaike, H.    

   
Akbay, P., Basaran, A.A., Undeger, U. & Basaran, N. 
 
dioica L. 
   
Bassett, I.J., Crompton, C.W. & Woodland, D.W. 

     
Berg, C.C.   

Böhle, U.-R., Hilger, H.H. & Martin, W.F. 
Echium
 
 
Bonsen, K.J. & Ter Welle, B.J.H.        

Boufford, D.E.  
        Flora of North Amer-
ica

Boufford, D.E. & Spongberg, S.A.       





Burger, W.     Flora
Costaricensis   
 
Chen, C.J.   Sphaerotylos 
Acta

Chen, C.J., Lin, Q., Friis, I., Wi lmot-Dear, C.M. & Monro, A.K.
      
Flora of China 

Chew, W.L. Laportea    
Singapore
Clarke, L.A., Rebelo, C.S., Goncalves, J., Boavida, M.G. & Jordan,
P. 

 
Collier, H.O.J. & Chesher, G.B.  
      
Chemotherapy
  
Conn, B.J. & Hadiah, J.T.   

    
Deng, T., Kim, C., Zhang, D.G., Zhang, J.W., Li, Z.M., Nie, Z.L. &
Sun, H.  Zhengyia shennongensis
     
  
  
    
    


Echium
Lobelia




 DendrocnideLaportea
Laportea
     
       


     
  

    
     
     

   Euphorbia  
   
        
   
 
       



        
      

    
     
Poikilospermum
Poikilospermum 



 Laportea
    
    

Laportea

ACKNOWLEDGEMENTS
      
      
Y.     
     
    


     
  


   
LITERATURE CITED
76
TAX O N
 Kim & al. • Phylogeny and character evolution of Urticeae
Version of Record
Felsenstein, J. 
Evolution
 
Friis, I.   Girardinia          

Friis, I.  Obetia      
   
Friis, I.   
     Evolution, systematics, and
.

Friis, I.   
 
   
and Caryophyllid families 
Geltman, D.V.        
  
Hadiah, J.T. & Conn, B.J. 
 Elatostema  
Blumea
Hadiah, J.T., Conn, B.J. & Quinn, C.J. 


Horn, J.W., Van Ee, B.W., Morawetz, J.J., Riina, R., Steinmann,
V.W., Berry, P.E. & Wurdack, K.J. 
Euphor-
bia 
   
Joly, S., Starr, J.R., Lewis, W.H. & Bruneau, A. 
 
Bot 
Kato, T., Ishida, K. & Sato, H.       

    
Kelchner, S.A.      
 

Knox, E.B., Downie, S.R . & Palmer, J.D. 
    Lobelias

Li, H.L. 

   
Maddison, W.P. & Maddison, D.R.        

Monro, A.K. 
    Pilea 

  
Monro, A.K. & Rodríguez, A.          

   
Niklas, K.J. 
Evolution
 
Posada, D. & Crandall, K.A.  
Bioinformatics
      
Qian, H.     
      

    
Rambaut, A. & Drummond, A.J. 
 
Ronquist, F. & Huelsenbeck , J.P.    
   Bioinformatics 
  
Soltis, D.E., Mort, M.E., Latv is, M., Mavrodiev, E.V., O’Meara,
B.C., Solt is, P.S., Burleigh, J.G. & De Casas, R.R. 
                 
 
 
Steinmann, V.W. 

Swofford, D.L. *
*

Sytsma, K.J., Morawetz, J., Pires, J.C., Nepokroeff, M., Conti, E.,
Zjhra, M., Hall, J.C. & Chase, M.W.  
               rbcL
trnLFndhF
   
Taberlet, P., Gielly, L., Pautou, G. & Bouvet, J.      
         

  
Takhtajan, A.L. 
   
   
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. &
Higgins, D.G.    


 
Tuberville, T.D., Dudley, P.G. & Pollard, A.J.   

Laportea canadensis. Oikos 
 
Wang, W.T. & Chen, C.J.  
 
Weddell, H.A .     
Prodromus systematis naturalis regni vegetabilis

 
White, T.J., Birns, T., Lee, S. & Taylor, J. 

  
   -
tions
    
Woodland, D.W.  
Evolution,


Woodland, D.W., Bassett, I. J. & Crompton, C.W. 
        

Wu, Z.Y., Monro, A.K., Milne, R.I., Wang, H., Yi, T.S., Liu, J. & Li,
D.Z. 
 

      
Zhang, S.D., Soltis, D.E., Yang, Y., Li, D.Z. & Yi, T.S. 
     

     
Zurawski, G. & Clegg, M.T. 


 
77
Kim & al. • Phylogeny and character evolution of Urticeae
TAX O N
 
Version of Record
Appendix 1.
  
TRIBESpecies,  voucher specimen          rbcL 

URTICEAE Dendrocnide basirotunda     Deng 408**  *;
Dendrocnide basirotunda        Dendrocnide meyeniana 
    Dendrocnide urentissima
   ** *; Discocnide mexicana 
Girardinia diversifolia      diversifolia        
  Girardinia diversifoliadiversifolia      Girardinia
diversifolia diversifolia       Girardinia diversifolia diversifolia 
 Girardinia diversifolia suborbiculata
      Girardinia diversifolia triloba 
        Girardinia diversifolia triloba
           Gyrotaenia myriocarpa   
       Hesperocnide sandwicensis      
    Hesperocnide sandwicensis           Hesperocnide
sandwicensis      Hesperocnide tenella
*  * *; Laportea alatipes       
  Laportea alatipes    
Laportea aestuans               Laportea aestuans
      Laportea bulbifera 
* **; Laportea bulbifera   
   Laportea canadensis         
Laportea cuspidata** *; Laportea cuspidata
      Laportea interrupta
    Laportea interrupta        
 Laportea lanceolata              
Laportea ruderalis  Laportea
ruderalisNanocnide japonica
    Nanocnide japonica  
Nanocnide lobata         Nanocnide lobata
Obetia aldabrensis  
Obetia pinnatifida       Obetia
radula       Obetia radula 
  Poikilospermum suaveolens                  
Poikilospermum suaveolens        **  *; Poikilospermum
sp. Poikilospermum sp. 
Poikilospermum sp. Urera baccifera
   Urera baccifera      
 Urera caracasana   Urera e lata
          Urera elata
   Urera fischeri        
Urera fischeri       Urera hypselodendron   
       Urera hypselodendron    
Urera hypselodendron 
Urera sansibarica  Urera sansibarica
Urera trinervis 
  Urtica dioica             Urtica
dioica     Urtica fissa
            Urt ica f iss a            *
**; Urtica laetevirenslaetevirens  **  *;
Urtica laetevirens  dentata  * * *; Urtica
laetevirens  dentata         Urtica laetevirens  dentata 
         Urtica mairei        *
* *; Urtica maire i*  * *; Urtica mairei 
        Urtica massaica          
Urtica massaica      Urtica massaica  
       Urtica massaica    
    Urtica massaicaUrtica massaica
    Urtica massaica 
       Urtica massaica        
Urtica massaicaUrtica taiwaniana 
   Urtica taiwaniana Urtica
thunbergiana    Urtica thunbergiana
     Urtica triangularis         *
* *; Urtica triangularis * *  *; Urtica triangu-
laris     Zhengyiia shennongensis  
     *   **; Zhengyiia shennongensis             
  **  *BOEHMERIEAEBoehmeria clidemioides    
 Boehmeria penduliflora            Boehmeria
sp.   EL ATOSTE MATEAE Elatostema ficoides  
        Elatostema stewardii     

TAX O N
 Kim & al. • Phylogeny and character evolution of Urticeae
Version of Record
  Pilea cavaleriei                 
Pilea plataniflora     * *   *; Pilea sinofasciata   
     Pilea sp.   
  FORSSKAOLEAEDroguetia iners                        
   Forsskaolea candida          PARIETAR-
IEAERousselia humilis              Rousselia humilis
           OUTGROUPS: MORACEA EFatoua villosa  
    Morus alba  * * *; Sorocea affinis   *
  * *; Trophis racemosa  * * *CANNABACEAECannabis sativa    
       Humulus lupulus         
 
Appendix 1.



a      
Urticeae
Dendrocnide basirotunda 3
Dendrocnide meyeniana 3
Dendrocnide urentissima 3
 3
Girardinia diversifolia  diversifolia 
Girardinia diversifolia  suborbiculata 
Girardinia diversifolia  triloba 
 
 
Laportea aestuans 3
Laportea cuspidata 3
Laportea interrupta 3
Laportea ruderalis 3
 3
Nanocnide lobata 3
Obetia aldabrensis 3
Obetia pinnatifida 3
Obetia radula 3
Poikilospermum _ 
Poikilospermum _ 
Poikilospermum suaveolens 
 3
 3
 3
 3
 3
 3
 3
 3
 3
Appendix 2.
a, habit,
b, leaf arrangement,c, number of stipules, 
d, stipule position,e, achene shape, f, bulbils,
g, stinging hairs,h, number of tepals, 

Urticeae 
 dentata 3
 laetevirens 3
 3
 3
 3
 3
 3
Zhengyia shennongensis 3
Elatostemateae
Gyrotaenia myriocarpa 
Elatostema ficoides        
        
Pilea cavaleriei 
Pilea plataniflora 
Pilea sinofasciata 
Pilea         
Boehmerieae
Boehmeria clidemioides 
Boehmeria penduliflora 
Boehmeria        
Parietarieae
      
Forsskaoleae
Droguetia iners 
Forsskaolea candida 
Outgroups
Morus alba 
Fatoua villosa 
Cannabis sativa 
 


a      
... This generic circumscription of the Urticeae, however, was established prior to the era of molecular phylogenetics. With the advent of the molecular tools, classification within tribe Urticeae has received much attention, with both taxonomic and phylogenetic studies spurring realignments (Hadiah et al., 2008;Kim et al., 2015;Huang et al., 2019;Wells et al., 2021). Molecular analyses have led to the exclusion of Gyrotaenia and the inclusion of Touchardia, Poikilospermum and Zhengyia in the tribe; hence, Urticeae presently comprises 12 genera (Wu et al., 2013;Kim et al., 2015;Jin et al., 2019). ...
... With the advent of the molecular tools, classification within tribe Urticeae has received much attention, with both taxonomic and phylogenetic studies spurring realignments (Hadiah et al., 2008;Kim et al., 2015;Huang et al., 2019;Wells et al., 2021). Molecular analyses have led to the exclusion of Gyrotaenia and the inclusion of Touchardia, Poikilospermum and Zhengyia in the tribe; hence, Urticeae presently comprises 12 genera (Wu et al., 2013;Kim et al., 2015;Jin et al., 2019). Molecular phylogenetic studies have also been able to demonstrate the monophyly of this tribe as well as which genera are polyphyletic or monophyletic. ...
... Wu et al. (2013), using seven combined markers from the mitochondrial, nuclear, and chloroplast genomes, recovered Laportea sister to a clade comprising Obetia-Urera-Touchardia and Poikilospermum, though with weak support ( Figure 1A). Subsequent studies, however, have supported alternative, conflicting resolutions of Laportea (Figures 1B-D; Kim et al., 2015;Wu et al., 2018;Huang et al., 2019) probably due to the limited sampling. The placement of Poikilospermum also remains uncertain; although it has consistently been placed sister to Urera, support for this was either lacking (Figures 1A-C; Wu et al., 2013Wu et al., , 2018Kim et al., 2015;Wells et al., 2021) or low ( Figure 1D; Huang et al., 2019). ...
Article
Full-text available
Urticeae s.l, a tribe of Urticaceae well-known for their stinging trichomes, consists of more than 10 genera and approximately 220 species. Relationships within this tribe remain poorly known due to the limited molecular and taxonomic sampling of previous studies, and chloroplast genome (CP genome/plastome) evolution is still largely unaddressed. To address these concerns, we used genome skimming data—CP genome and whole nuclear DNA (18S-ITS1-5.8S-ITS2-26S); 106 accessions—for the very first time to attempt resolving the recalcitrant relationships and to explore chloroplast structural evolution across the group. Furthermore, we assembled a taxon rich two-locus dataset of trnL-F spacer and nuclear ITS (nrITS) sequences across 291 accessions to complement our genome skimming dataset. We found that Urticeae plastomes exhibit the tetrad structure typical of angiosperms, with sizes ranging from 145 to 161 kb and encoding a set of 110 to 112 unique genes. The studied plastomes have also undergone several structural variations, including inverted repeat (IR) expansions and contractions, inversion of the trnN-GUU gene, losses of the rps19 gene, and the rpl2 intron, and the proliferation of multiple repeat types; 11 hypervariable regions were also identified. Our phylogenomic analyses largely resolved major relationships across tribe Urticeae, supporting the monophyly of the tribe and most of its genera except for Laportea, Urera, and Urtica, which were recovered as polyphyletic with strong support. Our analyses also resolved with strong support several previously contentious branches: (1) Girardinia as a sister to the Dendrocnide-Discocnide-Laportea-Nanocnide-Zhengyia-Urtica-Hesperocnide clade and (2) Poikilospermum as sister to the recently transcribed Urera sensu stricto. Analyses of the taxon-rich, two-locus dataset showed lower support but was largely congruent with results from the CP genome and whole nuclear DNA dataset. Collectively, our study highlights the power of genome skimming data to ameliorate phylogenetic resolution and provides new insights into phylogenetic relationships and chloroplast structural evolution in Urticeae.
... Stinging trichomes are structures typically associated with species of Urticaceae [6], which is the only family in the order Rosales with this type of trichome. More precisely, these trichomes occur only in Urticeae and may be a synapomorphy of the tribe, which has been hypothesized to promote species diversification [50]. Urticeae have 12 genera, of which 11 have stinging trichomes ( Table 2). ...
... The possibility is reinforced by the fact that the genus Gyrotaenia had been originally described as belonging to tribe Urticeae and having stinging trichomes [5]; however, Kim et al. [34] describe the genus as having no stinging trichomes. It has recently been proposed that Gyrotaenia is closer to tribe Elatostemateae than to Urticeae, considering the absence of stinging trichomes and the occurrence of female flowers with two tepals [40,50]. Both characters are important for the circumscription of Urticeae [50]. ...
... It has recently been proposed that Gyrotaenia is closer to tribe Elatostemateae than to Urticeae, considering the absence of stinging trichomes and the occurrence of female flowers with two tepals [40,50]. Both characters are important for the circumscription of Urticeae [50]. Our analysis indicates that the stinging trichomes evolved once in Urticaceae. ...
Article
Full-text available
Stinging trichomes are rare in plants, occurring only in angiosperms, where they are reported for a few genera belonging to six families. Although there is no report of stinging trichomes in Apocynaceae, previous fieldwork collections of Fischeria and Matelea caused us a mild allergic reaction on the skin when we contacted the dense indumentum of the plants. This fact associated with the well-known presence of glandular trichomes with acute apex in both genera raised suspicions that stinging trichomes could be present in the family. Hence, this study aimed to investigate the likely occurrence of stinging trichomes in Fischeria and Matelea. We analyzed vegetative shoots and leaves of Fischeria stellata and Matelea denticulata through the usual procedures of light and scanning electron microscopy. We also performed several histochemical tests to investigate the chemical composition of trichome secretion. We detected that glandular trichomes occur throughout the surface of the leaf and stem. They are multicellular, uniseriate with an apical secretory cell, which has a dilated base and a needle-shaped apex. The secretion is compressed into the acuminate portion of the apical cell by a large vacuole, and crystals are deposited in the cell wall in a subapical position, providing a preferential site of rupture. The secretion, composed of amino acids and/or proteins, is released under mechanical action, causing skin irritation. Based on our detailed morphological and anatomical analyses, and in the functional aspects observed, we concluded that the glandular trichomes in Fischeria and Matelea can indeed be classified as stinging. Thus, Apocynaceae is the seventh family for which this type of trichome has been reported. We also compiled information on stinging trichomes in all families of angiosperms. Their phylogenetic distribution indicates that they have evolved at least 12 times during angiosperm evolution and may represent an evolutionary convergence of plant defense against herbivory.
... The nettle family Urticaceae comprises 53 genera grouped into four tribes and encompassing c. 2150 species of herb, shrub, tree and vine distributed throughout the tropical and temperate regions of the world. Of the four tribes, the Urticeae contains c. 10 genera (Friis, 1989a) , among which both Wu et al. (2013) and Kim et al. (2015) recovered extensive polyphyly focussed around the genus Urera Gaudich.. Urera was first described by Gaudichaud-Beaupré (1830) to account for several specimens of Urtica-like plants from the Neotropics and South East Asia that bore fruits whose perianths become fleshy upon ripening, and for which Urera baccifera (L.) Gaudich. ex Wedd. ...
... In the phylogenetic analyses of Kim et al. (2015) and Wu et al. (2013), three currently recognised genera were found to be nested within the traditional circumscription of Urera: the African xerophytic trees and shrubs, Obetia Gaudich. (7 spp.), the Hawaiian tree, Touchardia Gaudich. ...
... We expanded the taxon and genome sampling used for the phylogenies of Wu et al. (2013) and Kim et al. (2015), and further combined the results with a detailed review of the morphology. We applied a two-tier genome sampling approach, where (i) a taxon-dense sample of Sanger sequence data from one nuclear and one chloroplast barcoding region was used to recover monophyletic clades of putative generic rank, and (ii) a genome-dense sample of target-enrichment data for a subsample of representatives from each well-supported clade from the Sanger data was used to resolve relationships among them. ...
Article
Urera Gaudich, s.l. is a pantropical genus comprising c. 35 species of trees, shrubs, and vines. It has a long history of taxonomic uncertainty, and is repeatedly recovered as polyphyletic within a poorly resolved complex of genera in the Urticeae tribe of the nettle family (Urticaceae). To provide generic delimitations concordant with evolutionary history, we use increased taxonomic and genomic sampling to investigate phylogenetic relationships among Urera and associated genera. A cost-effective two-tier genome-sampling approach provides good phylogenetic resolution by using (i) a taxon-dense sample of Sanger sequence data from two barcoding regions to recover clades of putative generic rank, and (ii) a genome-dense sample of target-enrichment data for a subset of representative species from each well-supported clade to resolve relationships among them. The results confirm the polyphyly of Urera s.l. with respect to the morphologically distinct genera Obetia, Poikilospermum and Touchardia. Afrotropic members of Urera s.l. are recovered in a clade sister to the xerophytic African shrubs Obetia; and Hawaiian ones with Touchardia, also from Hawaii. Combined with distinctive morphological differences between Neotropical and African members of Urera s.l., these results lead us to resurrect the previously synonymised name Scepocarpus Wedd. for the latter. The new species epiphet Touchardia oahuensis T.Wells & A.K. Monro is offered as a replacement name for Touchardia glabra non H.St.John, and subgenera are created within Urera s.s. to account for the two morphologically distinct Neotropical clades. This new classification minimises taxonomic and nomenclatural disruption, while more accurately reflecting evolutionary relationships within the group.
... We focussed on the ITS nuclear region and the rbcL and trnL-F plastid regions. We combined sequences generated by previous studies (Monro, 2006;Jestrow & al., 2012;Wu & al., 2013Wu & al., , 2018Kim & al., 2015) with 159 sequences generated by ourselves. We excluded accessions when we felt that the identifications were ambiguous, or sequence data for only a single region could be obtained. ...
... Lecanthoides (all of the species). Four species of Moraceae (Fatoua villosa Nakai, Morus alba L., Sorocea affinis Hemsl., Trophis racemosa (L.) Urb.), two species of Cannabaceae (Cannabis sabiva L., Humulus lupulus L.) and 13 species from Urticaceae were chosen as outgroups based on previous analyses (Zhang & al., 2011;Kim & al., 2015). Species names, the accession numbers of sequences downloaded from GenBank, and newly generated sequences used in this study are listed in Appendix 1. ...
Article
Full-text available
Ensuring that genera are monophyletic should result in a stable classification that better reflects evolutionary history, and which can serve as a framework for answering broader scientific questions. It is also important that species can be assigned to genera easily and unambiguously so as to remove barriers to identification and species discovery. Pilea, with 933 published names and likely more than 700 spp. is the largest genus within the Urticaceae and was last monographed in 1869. Whilst the monophyly of the genus has been proposed by previous authors, this has been based on incomplete taxon sampling that has failed to resolve the position of key taxa. We aimed to generate a robust phylogeny for Pilea and allied genera that could test the monophyly of the genus and provide a framework for a taxonomic revision of this species‐rich yet poorly studied genus. To do so, we sought to sample taxa encompassing previous infrageneric classifications and species with anomalous inflorescences or flower morphology and to use the resulting phylogeny to evaluate the delimitation of Pilea and to establish an infrageneric classification. In addition, we included a representative of the Polynesian genus Haroldiella, which, morphologically, seemed indistinguishable from Pilea. Using two plastid and one nuclear region, we constructed phylogenies using Bayesian inference, maximum likelihood and maximum parsimony approaches. We used our BI phylogeny to evaluate the informativeness of 22 morphological characters and used both to delimit monophyletic groups and infrageneric sections. Our results recovered Pilea as paraphyletic, a consequence of the recovery of a monophyletic clade comprising Lecanthus and Pilea sect. Achudemia and sect. Smithiella, neither of which had been adequately sampled in previous studies. We also recovered Pilea as paraphyletic with respect to Haroldiella. Pilea (including Haroldiella) was recovered as sister to the clade comprising Lecanthus and Pilea sect. Achudemia and sect. Smithiella. We identified isomery between male and female flowers, flower part number, male sepal arrangement and achene ornamentation as being phylogenetically informative characters that can be used to distinguish Achudemia (Pilea sect. Achudemia and sect. Smithiella) from Pilea. We also identified a trend of decreasing merism and fruit size, correlated with an increase in species‐richness across the phylogeny which our results suggest may indicate a shift in pollen and fruit dispersal, pollen dispersal becoming more kinetic and fruit dispersal less reliant on animals. On the basis of our evaluation of both morphological characters and phylogenetic relationships we propose a new infrageneric classification for the genus comprising eight sections, five of which we propose for the first time here: Pilea sect. Angulatae, sect. Plataniflorae, sect. Tetraphyllae, sect. Trimeris and sect. Verrucosae.
... Morphological study of G. diversifolia showed high level of variation mostly on its leaf lobes [6,8,14]. Leaves are alternate, petiolate, and elliptic to ovate, but consist of varying degrees of division in the same plant or within the population [2,6,8,15,16]. Genetic analysis of plants provides broader knowledge on their diversity and basis to study important metabolites that they produce [17]. ...
Article
Full-text available
Girardinia diversifolia (Link) Friis from Urticaceae family is a perennial herb, commonly known as the Himalayan Giant Nettle. It has cultural, medicinal, and economic values among the indigenous peoples and local communities living in the mountains of Nepal and India. Prior knowledge of genetic diversity in plant species may contribute to conservation and sustainable utilization of important genotype. The aim of this study is to assess the genetic diversity within and among different populations of the 45 accessions of G. diversifolia collected from Far-western, Central, and Eastern regions of Nepal. The amplification of genomic deoxyribonucleic acid (DNA) with ten inter-simple sequence repeat (ISSR) primers yielded 131 clear DNA bands, of which 98.09% were found to be polymorphic. The mean effective number of alleles (ne), Nei's gene diversity (h) and Shannon's information index (I) were 1.598, 0.349, and 0.523, respectively. The total genetic diversity Ht (0.350 ± 0.04), low intrapopulation genetic diversity Hs (0.141 ± 0.03) and low estimated gene flow Nm (0.355 ± 0.11) reflected high genetic differentiation among population Gst (0.594 ± 0.08). The analysis of molecular variance among and within five populations of G. diversifolia showed that the value of genetic variation among the population was 60% whereas the value of genetic variation within the population was 40%. The results show that the ISSR markers are informative for the study of genetic diversity, which may help in the conservation and sustainable utilization of G. diversifolia and allied taxa.
... Family Urticaceae comprises 54 genera and more than 2000 species of herbs, shrubs, small trees, and a few vines, distributed primarily in tropical regions [1,2]. A survey of literature on family Urticaceae showed many reports about antidiabetic, anti-inflammatory and antimicrobial activities [3]. ...