A classification of Selaginella (Selaginellaceae) based on molecular (chloroplast and nuclear), macromorphological, and spore features

Abstract

The largest genus of seed-free vascular plants Selaginella alone constitutes the family Selaginellaceae, the largest of the lycophyte families. The genus is estimated to contain ca. 800 species distributed on all continents except Antarctica, with the highest species diversity in tropical and subtropical regions. The monophyly of Selaginella has rarely been doubted, in contrast its infrageneric classification has been contentious. In the present study, based on chloroplast and nuclear DNA evidence, macromorphology, spore features, and/or distribution information, Selaginella is classified into six subgenera: S. subg. Selaginella, S. subg. Boreoselaginella, S. subg. Pulviniella, S. subg. Ericetorum, S. subg. Heterostachys, and S. subg. Stachygynandrum. The latter three subgenera are further classified into six, five, and seven sections, respectively. All of these infrageneric divisions, identified with molecular data, are supported by non-molecular features. A key to infrageneric taxa is given. Thirty-seven infrageneric taxa published in earlier literature are lectotypified and classified into those infrageneric taxa here recognized. A nomenclatural account of each infrageneric taxon is given.

Full text

1117
Zhou & Zhang • Classification of Selaginella
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–
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
n/a;  || 
INTRODUCTION
Selagi-
nella

 



 

   
 



Selaginella moellendorffii 
        
  
   
Selaginella


   
Selaginella  
     
   
& 

Selaginella 



S. Tetragonostachys




         
   


A classification of Selaginella (Selaginellaceae) based on molecular
(chloroplast and nuclear), macromorphological, and spore features
Xin-Mao Zhou1,2 & Li-Bing Zhang3
 Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization, Chengdu Institute of Biology, Chinese Academy
of Sciences, P.O. Box 416, Chengdu, Sichuan 610041, China
 School of Life Sciences, Sichuan University, Chengdu, Sichuan 610064, China
 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.
 Li-Bing Zhang, Libing.Zhang@mobot.org

DOI
 
Abstract
   Selaginella
     

  


 

 Selaginella 

Selaginella
S. SelaginellaS. BoreoselaginellaS. PulviniellaS. EricetorumS. Heterostachys  S. 
Stachygynandrum
 


Keywords
Selaginella
Supplementary Material


SYSTEMATICS AND PHYLOGENY

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–Zhou & Zhang • Classification of Selaginella
Version of Record
Fig. 1.
   Selaginella I.
A,
S  Selaginella;
B,
SBoreoselaginella;
C–I,
S  Ericetorum;
J,
S
Pulviniella;
K–P,
SHeterostachys
A,
Selaginella deflexa S  Selaginella
B,
S sanguinolenta SBoreoselaginella
C,
S uli-
ginosa S.  Lyallia
D,
S scopulorum S. Homoeophyllae
E,
S lepidophylla S. Lepidophyllae
F,
S remotifolia S. Articu-
latae
G,
S diffusa S. Articulatae
H,
S exaltataS. Megalosporarum
I,
S myosurus S. Myosurus
J,
S pulvinata S
Pulviniella
K
S douglasii S. Auriculatae
L,
S nipponica S. Homostachys
M,
S braunii S. Oligomacrosporangiatae
N,
S uncinata S. Oligomacrosporangiatae
O,
S delicatula S. Oligomacrosporangiatae
P,
S heterostachys S. Heterostachys
 


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Zhou & Zhang • Classification of Selaginella
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INFRAGENERIC TAXONOMIC HISTORY
Selaginella
S. selaginoides
          

Selag-
inella  Lycopodium 
Selaginella


  

 

Selaginella
Lycopodium
Fig. 2.
        Selaginella 
A–B,
S  Heterostachys;
C–N,
S  Stachygynandrum
A,
Selaginella bodinieri
S. Heterostachys
B,
S compta S. Tetragonostachyae
C–D,
S biformis S. Plagiophyllae
E,
S involvensS. Circinatae
F,
S moellendorffii S. Circinatae
G,
S oaxacana S. Proceres
H,
S apoda S. Pallescentes
I,
S pallescensS. Palle-
scentes
J,
S novae-hollandiae S.  Austroamericanae
K,
S flagellata S. Heterophyllae
L,
S haematode S. Heterophyllae
M,
S doederleinii S. Ascendentes
N,
S frondosa S. Ascendentes— –

1120
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Fig. 3.
Selaginella 
A,
S  Selaginella;
B,
SBoreoselaginella;
C–I,
S  Ericetorum;
J,
SPulviniella;
K–P,
SHeterostachys
A,
S. deflexa S  Selaginella
B,
S sanguinolenta SBoreoselaginella
C,
S uliginosa S.  Lyallia
D,
S scopulorum S. Homoeophyllae
E,
S lepidophylla S. Lepidophyllae
F,
S remotifolia S. Articulatae
G,
S diffusa S. 
Articulatae
H,
S. exaltataS. Megalosporarum
I,
S myosurus S. Myosurus
J,
S pulvinata SPulviniella
K,
S douglasii
S. Auriculatae
L,
S nipponica S. Diplostachyum
M,
S braunii S. Oligomacrosporangiatae
N,
S uncinata S. Oligo-
macrosporangiatae
O,
S delicatula S. Oligomacrosporangiatae
P,
S labordei S. Heterostachys. —       
 

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Zhou & Zhang • Classification of Selaginella
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L. Exstipulatae 
L. Stipulatae 
Lycopodium Selaginella. 
   
SelaginellaL. 
Spicata    L.  Stachygynan-
drum        L. 
Circinata  L.  Planifolia 
L.      Tetragonostachya   
 L.  Platy stachya    
  
     
 Selaginella
         Lycopodium
Fig. 4.
Selaginella
A–B,
S  Heterostachys;
C–L,
S  Stachygynandrum
A,
Selaginella bodinieri S. Hete-
rostachys
B,
S compta S. Tetragonostachyae
C,
S biformis S. Plagiophyllae
D,
S involvensS. Circinatae
E,
S moellen-
dorffi S.  Circinatae
F,
S oaxacana S.  Proceres
G,
S apoda S.   Pallescentes
H,
S pallescensS.  Pallescentes
I,
S novae-hollandiae S.  Austroamericanae
J,
S flagellata S. Heterophyllae
K,
S haematodes S. Heterophyllae
L,
S doe-
derleinii S. Ascendentes–

1122
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Fig. 5.
Selaginella
A,
S.Boreoselaginella;
B, C, N, O,
S.  Ericetorum;
D, P,
S.Pulviniella;
E–J, Q–V,
S.Heterostachys;
K, L, W, X,
S. Stachygynandrum
A–L,

M–X,

A, M,
S. san-
guinolenta S.Boreoselaginella
B, N,
S indica S. 
Homoeophyllae
;
C, O,
S. remotifolia S. 
Articulatae
;
D,P,
S. stauntoniana
S.Pulviniella
E, Q,
S. heterostachys S. Heterostachys
F, R,
S. ciliaris S. Tetragonostachyae
G, S,
S. pallidissima S. 
Homostachys
H, T,
S. braunii S. 
Oligomacrosporangiatae

I, U,
S. pseudopaleifera S. 
Oligomacrosporangiatae

J, V,
S. pennata
S. 
Oligomacrosporangiatae

K, W,
S. involvens S. Circinatae
L, X,
S. doederleinii S. 
Ascendentes


1123
Zhou & Zhang • Classification of Selaginella
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–
Version of Record
  
Selaginella    
  S.Selaginella 
 S.Homoeophyllae S.
Hetero phyllae S.Dip-
lostachyumS.Stachygynandrum
   
 S.Dicho tropae 
S. Homotropae   
S.Dichotropae
 S.  Homotropae  
S.
SelaginellaS. Homostachys  S. Hetero-
stachys S.Stachy gynandrum 
Sela-
ginella S.Homoeo phyllum
S.Heterophyllum 
 

S.Boreoselaginella
S. Selaginella
“S.Euselaginella” S. Selaginella
 
 “S.Euselaginella
S. Selaginella
 S.SelaginellaS. Ericeto-
rum S.Heterostachys S.Tetragonostachys
S. Stachygynandrum
    








MATERIALS AND METHODS


     


 

   



 
 Melbourne Code



   
   Seleginella    
 

  

 
  




  


 

 


 



Zhou &   
 
Selaginella




RESU LT S
Selaginella
 
Selaginella
  

Selaginella  




  
 

  
 




 



1124
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–Zhou & Zhang • Classification of Selaginella
Version of Record
Tab le 1.
Selaginella 


   a

S. Selaginella Homoeophyllum
Cylindrostachyae Euselaginella Selaginella Selaginella

S. Boreoselaginella Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Stachygynandrum
Decumbentes Stachygynandrum Lycopodiodes

S. Ericetorum Homoeophyllum
Heterophyllum Euselaginella
Stachygynandrum Ericetorum
Stachygynandrum 
Tetragonostachys
Lycopodiodes

S. Articulatae Heterophyllum
Oligomacrosporangiatae
Articulatae
Stachygynandrum
Articulatae Stachygynandrum Lycopodiodes

S. Homoeophyllae Homoeophyllum
Tetragonostachyae Euselaginella Tetragonostachys Bryodesma

S. Lepidophyllae Heterophyllum
Pleiomacrosporangiatae Stachygynandrum Stachygynandrum Lycopodiodes

S. Lyallia Homoeophyllum
 Tetragonostachyae
Heterophyllum
Pleiostelicae
Euselaginella 
Stachygynandrum  Ericetorum 
Stachygynandrum Lycopodiodes

S. Megalosporarum Heterophyllum
Oligomacrosporangiatae
Articulatae
Stachygynandrum
Articulatae Stachygynandrum Lycopodiodes

S. Myosurus Homoeophyllum Stachygynandrum Stachygynandrum Lycopodiodes

S. Pulviniella Heterophyllum
Pleiomacrosporangiatae Stachygynandrum
Circinatae Stachygynandrum Lycopodiodes

S. Heterostachys Heterophyllum  Heterostachys
 Stachygynandrum 
Homostachys
Heterostachys
Stachygynandrum Lycopodiodes

S. Auriculatae Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Stachygynandrum Stachygynandrum Lycopodiodes

S. Heterostachys Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Heterostachys Heterostachys Lycopodiodes

S. Homostachys Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Homostachys Stachygynandrum Lycopodiodes

S. Oligomacro-
sporangiatae
Heterophyllum
Pleiomacrosporangiatae
Pleiostelicae 
Oligomacrosporangiatae
Monostelicae
Stachygynandrum
Sarmentosae
Caulescentes
Circinatae
Heterostachys
Stachygynandrum
Heterostachys Lycopodiodes

Tetragonostachyae Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Heterostachys Heterostachys Lycopodiodes

S. Stachygynandrum Heterophyllum Stachygynandrum 
Heterostachys Stachygynandrum 
Heterostachys Lycopodiodes

S. Ascendentes Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Stachygynandrum Stachygynandrum Lycopodiodes

S. Austroamericanae Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Heterostachys Heterostachys Lycopodiodes


Zhou & Zhang • Classification of Selaginella
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–
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DISCUSSION AND TAXONOMIC
TREATMENT
          
Selaginella
  
 



Selaginella LycopodioidesLycopodium
Stachygynandrum
SelaginellaDidiclisLyco-
podioidesSelaginella  


Selaginella  S.  Selaginella Lycopodioi-
desS.  Ericetorum    +S.Heterostachys
S.  Tetragonostachys
Bryodesma   Tetragonostachys 
Selaginella
Bryodesma  
BryodesmaLycopodioides

    
Selagi-
nella
Selaginella   [
  Stachygynandrum    
& 
Lycopodium selaginoides Selaginella selaginoides

Selaginoides 
Lycopodium selaginoides 
Mirmau
Lycopodium selaginoides
Polycocca  
Lycopodium selaginoides
Trispermium
Lycopodium selaginoides
Lycopodioides   
    
   
Lycopodioides denticulata
Selaginella denticulata 
Didiclis & 
Didiclis ornithopodioides
Selaginella ornithopodioides

Gymnogynum
Didiclis Gymnogynum
Gym-
nogynum domingenseSelaginella plumosa  

Diplostachyum
Diplostachium Lycopodium  Diplostachyum
Selaginella
Diplostachyum
Lycopodioides Diplostachyum 

Diplostachyum tenellum Selagi-
nella tenella 
Hypopterygiopsis     
Hypopterygiopsis reptansSelaginella
sakuraii 
Tab le 1.



   a

S. Circinatae Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Stachygynandrum Stachygynandrum Lycopodiodes

S. Heterophyllae Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Stachygynandrum
Heterostachys Stachygynandrum
Heterostachys Lycopodiodes

S. Pallescentes Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Stachygynandrum  Stachygynandrum Lycopodiodes

S. Plagiophyllae Heterophyllum
Pleiomacrosporangiatae
Monostelicae
Stachygynandrum Stachygynandrum Lycopodiodes

S. Proceres Heterophyllum
 Pleiomacrosporangiatae
Monostelicae
Stachygynandrum
Heterostachys
Stachygynandrum
HeterostachysLycopodiodes
a 1993Selaginella Bryodesma Lycopodiodes


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–Zhou & Zhang • Classification of Selaginella
Version of Record
Fig. 6.
Selaginella rbcL


0.05 S. siamensis subclade
Sect. Megalosporarum
Sect. Myosurus
Sect. Plagiophyllae (Asia)
Pacific
Asia
Sect. Auriculatae
S. braunii subclade
S. willdenowii subclade
S. delicatula
subclade
S. pervillei subclade
S. pennata subclade
S. stauntoniana subclade
S. imbricata subclade
Sect. Circinatae (Asia)
Sect. Heterophyllae (America)
Sect. Ascendentes (Asia+Australia)
Sect. Lyallia
Sect. Heterostachys
Sect. Homostachys
Sect. Lepidophyllae
Sect. Homoeophyllae
Sect.
Oligomacro-
sporangiatae
Outgroups
Sect. Tetragonostachyae
Sect. Pallescentes (America)
Sect. Poceres (America)
Sect. Austroamericanae (America)
100/100/1.00
100/100/1.00
91/96/0.99
91/88/0.89
45/*/0.74
99/100/1.00
60/55/*
62/61/*
94/97/1.00
79/69/0.97
81/85/1.00
95/87/1.00
69/82/0.99
100/100/0.99
99/99/0.99
100/100/1.00
82/74/0.97
94/96/1.00
90/93/0.95
73/51/0.78
100/100/1.00
100/100/1.00
100/99/0.99
97/97/0.99
80/73/0.99
81/82/1.00
90/97/1.00
100/100/1.00
100/91/0.99
99/98/1.00
100/100/100
100/100/1.00
100/100/1.00
55/81/0.94
100/100/1.00
100/100/0.99
98/99/0.99
100/98/1.00
89/*/0.99
99/*/0.99
100/100/1.00
Subg.
Stachygynandrum
Subg.
Heterostachys
Subg.
Ericetorum
Subg.
Pulviniella
Subg.
Boreoselaginella
Subg.
Selaginella
*/53/0.99
*/78/1.00
100/100/1.00
55/61/0.99
100/100/1.00
Sect. Articulatae

1127
Zhou & Zhang • Classification of Selaginella
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–
Version of Record
Bryodesma  
Bryodesma rupestre Selaginella rupestris

Lycopodium selaginoides 

   
Selaginella spinosa
Selaginella
S. spinosa L. selaginoides
  
Lycopodioides
Lycopodioides denticulata 


 

SelaginellaSelaginella
SelaginellaS. Boreoselaginella S. Eri-
cetorum  S. Heterostachys S. Stachy-
gynandrum   S. Pulviniella 

  

 S. EricetorumS. Heter-
ostachysS. Stachygynandrum
 

Selaginella 
Selaginella 
S. Selaginella
S. Boreoselaginella 
S. Ericetorum 
S. 
Articulatae

& 
S. 
Homoeophyllae

S. Lepidophyllae

S. 
Lyallia (


S. Megalosporarum 

S. Myosurus

S. Pulviniella 

S. Heterostachys 
S. Auriculatae

S. Heterostachys 

S. Homostachys 

S. 
Oligomacrosporangiatae


S. Tetragonostachyae 

S. Stachygynandrum 

S. 
Ascendentes


S. Austroamericanae 

S. Circinatae 

S. Heterophyllae 
S. Pallescentes

S. Plagiophyllae 

S. Proceres 

Key to subgenera and sections (see Table 2)
 
n 
 S.  Selaginella
 

n
  
      
S. rossii

 
 S.  Boreoselaginella
 


S.Lepidophyllae
   



         

  


S.  Lyallia S.  Ericetorum 
  
 
 
 
 S.  Lepidophyllae
 

     

n
 S.  Homoeophyllae
 
 

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–Zhou & Zhang • Classification of Selaginella
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 
 
n S.  Lyallia
       
 
 n
 S.  Myosurus
  
 
 S.  Megalosporarum
  

n S.  Articulatae
 

S. stauntoniana


n S.  Pulviniella
 

     
S. pallescens
 
      





S.Tetragonostachyae
 S.  Heterostachys 
       

 S.  Stachygynandrum
         
       
S. bisulcata S. pennata 
 

S. denticulataS. helvetica 
 



n  S.  Oligomacrosporangiatae
 

 S.  Auriculatae
  

n 
 S.  Homostachys
 

 
 
   
n
 S.  Heterostachys
 

 S.  Tetragonostachyae
  
n
S.  Plagiophyllae
  
 
n S.  Ascendentes
  
 
  
n S.  Circinatae
        
 
 
 
 S. apoda




S. oaxacana 
    
     

  


   
 
n
 S.  Pallescentes
   
n
 S.  Proceres
   
 S.  Austroamericanae
 
 n
 S.  Heterophyllae
SelaginellaSelaginella

Lycopodium

Exstipulata


Exstipulatae
Lectotype (desig-
nated here):
Lycopodium selaginoides


Lycopodium

Spicata
 
Lectotype (designated here):
Lycopodium
selaginoides

Selaginella selaginoides



Selaginella

Homoeophyllum
  

Selaginella selaginoides  
.
Selaginella Homotropae 
        Lectotype

1129
Zhou & Zhang • Classification of Selaginella
TAX ON
–
Version of Record
(designated here): Selaginella spinulosa  
Selaginella selaginoides 
Selaginella Cylindrostachyae
S.  Cylindro-
stachyae
 Selaginella spi-
nulosa 
Selaginella  Polystichae 
 – Lectotype (designated
here): Selaginella spinulosa 
Selaginella spinulosa 
S. Cylindrostachyae 
Selaginella spi-
nulosa 
 S.  Polystichae  S.
Homotropae
 
Lycopodium
selaginoides


L.

Spicata

L.

Exstipulata


“Selaginella  Euselaginella   
“S.Euselaginella


 Selaginella Homotropae

S.

Homoeophyllae



SelaginellaSelaginella 
    


S. selaginoidesn

SelaginellaSelaginella
S. selaginoides
  S. deflexa  



S. Homoeophyllae  S.
Homotropae S. Cylindrostachyae
S.Euse-
laginella    
S. Selaginella

S. Selaginella    


 

SelaginellaBoreoselaginella.
Lectotype (designated here): Selagi-
nella borealis  Selaginella sanguinolenta

   
Selaginella borealis  

Selagi-
nella Boreoselaginella   
S. sanguinolenta


  

 S. sanguinolenta
 & S. nummularifolia


SelaginellaEricetorum   
S. uliginosa
Selaginella  Tetragonostachys

Selaginella rupestris 
        

  



Lycopodium
 TetragonostachyaSelaginella
 Tetragonostachyae 
   S. 
Tetragonostachys 


   
  
S.  Tetragonostachyae 
S. Heterostachys 
Selaginella  Ericetorum

    S. exaltata S. lepidophylla S. myosu-
rusS.Ericetorum  S. Articulatae
S. Tetragono-
stachys          S. lyalii
  S. moratii   
S. polymorpha 
  
Selaginella Tetragonostachys  
S.
Tetragonostachys
S. Lyallia
 

S. lepidophylla


        


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–Zhou & Zhang • Classification of Selaginella
Version of Record
Tab le 2.
Selaginella 
   

  

 

S.Selaginella   

 


 18 


S. Boreoselaginella   



 




 

S.Ericetorum  





  




S. Articulatae   

   20 



S. Homoeophyllae   

 


 18 





S. Lepidophyllae  




 







 

S. Lyallia  





 







 18 

S. Megalosporarum 

 

    

S. Myosurus   

  
 18–20 

S. Heterostachys 


 













S. Auriculatae   

 



 

1131
Zhou & Zhang • Classification of Selaginella
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–
Version of Record

S. Heterostachys 


 







 

S. Homostachys 

 










 18 

S. Oligomacrosporangiatae 


 













 



Tetragonostachyae 

 



 

 


S. Pulviniella   

 





20 


S. Stachygynandrum 


 











S. Ascendentes 

 

 



 18 

S. Austroamericanae 

 



   

S. Circinatae   

  
 20 

S. Heterophyllae 

 




  20 

S. Pallescentes 

 

    

S. Plagiophyllae 


 

   20 

S. Proceres 

 

    

1132
TAX ON
–Zhou & Zhang • Classification of Selaginella
Version of Record
 Selaginella
Articulatae
 
&  
Selaginella

Articulatae


S.

Articulatae


Didiclis

Articu-
latae

 
 
Didiclis sulcata
 
Selaginella
sulcata

= Selaginella

Adscendentes

 –
Lecto type (designated here):
Selaginella galeottii

 S. stellata 
= Selaginella

Caulescentes

–

Selaginella asperula

= Selaginella
  
Repentes
   
–
Lecto type (designated here):
Selaginella hortensis

 S. kraussiana
= Selaginella

Monostelicae

 
& & 

Selaginella remotifolia

= Selaginella

Pleiostelicae

  
&   &   
   Selaginella kraussiana    

.
Selaginella Arti-
cu latae 
ICN & 
Gymnogynum domingense 
Gymnogynum 

Didiclis stolonifera 


S. plumosaS
Articulatae
Didiclis Articulatae D. 
Gymnogynum  


Selaginella articulata
  
  

 Selaginella
sulcata


Selaginella
galeottii


S.

Adscendentes
 


Selaginella

Repentes

          


S. helvetica

S. hortensis
 
 
S.

Articulatae

S.

Homostachys


  
Selaginella

Adscendentes
    
  

 
S. gale-
ottii

S. flexuosa


S.

Articulatae

S.

Heterophyllae




Selaginella asperula


S.

Caulescentes


S.

Articulatae.
Selaginella Articulatae  

S. Articulatae


 

  







 Selaginella 
Homoeophyllae

Selaginella Homotropae

–

Selaginella rupestris

Selaginella rupestris   S. 
Homoeophyllae
 
 
S. spinosa S. selaginoides 
S. rupestrisS. 
Homoeophyllae
S. 
Homoeophyllae


Selaginella  Homoeophyllae S.
Tetragonostachys              




 



  

  

1133
Zhou & Zhang • Classification of Selaginella
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–
Version of Record


   


    Selaginella  Lepidophyllae
&  sect. nov.Selaginella lepidophylla




Selaginella Lepidophyllae
S. lepidophylla

  

  

 

S. novoleonensisS. ribae 
SLepidophyllae

 SelaginellaLyallia 
& comb. nov.DidiclisLyallia

Selaginella lyallii


Selaginella

Pleiostelicae


& & 
    – Lectotype (designated here):
Selaginella
lyallii

= Selaginella

Tetrastichae

– Lectotype (designated
here):
Selaginella uliginosa


Selaginella

Ericetorum


Selaginella uliginosa

  S
elaginella
 Ericeto-
rum  






  S. pygmaea
    S. moratii Rauh
 S. polymorpha  


n
S. uliginosaS. lyallii

        
    S
elaginella
aboriginalis 
  S. gracillima   S. uligi-
nosaS. royenii 
S. lyallii S. moratii S. polymorpha S. pygmaea
 

 Selaginella Megalosporarum 
&  sect. nov.   S
elaginella
exaltata



 
  Selaginella exaltata Selaginella exaltata 
   



  Selaginella  Myosurus
sect. nov.   S
elaginella
myosurus


   

Selaginella myosurus   

 S. scandens  S. puberula

 Selaginella  Pulviniella  
& subg. nov. S
elaginella
pulvinata




 Selaginella Rosulatae

S. Circinatae 









  S
elaginella
stauntoniana

S. imbricata
SelaginellaHeterostachys 

S
elaginella
heterostachys

S
elaginella
heterostachys 

1134
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–Zhou & Zhang • Classification of Selaginella
Version of Record
S. heterostachys

S. heterostachys

   Selaginella  Hetero-
stachys

S.Heterostachys
S. Heterostachys
S. Homostachys 
S. Suberosae
  
 S. douglasii   



S. brauniiS. delicatula S. hoffmannii S. pervil-
lei S. siamensisS. uncinataS. douglasii




 

S
elaginella
Stachygynandrum



          

  Selaginella Auriculatae
sect. nov.S
elaginella
douglasii 



    
Selaginella douglasii



  S. douglasii 
       S. Pleiomacrosporan-
giataeMonostelicae
S. delicatissima S. saccharata
  S. reflexa       
S. douglasiiS. delicatissima
  Selaginella Heterostachys   
& sect. nov. S
elaginella
heterostachys

LycopodiumPlatystachya
S
elaginella
Platystachyae

Lectotype (designated here): S
elaginella
chrysocaulos 
= Selaginella

Brachystachyae
 

Selaginella brachystachya


.


  


S. Tetragonostachyae

 


Selaginella brachy-
stachyaS. daozhenensis 

  
 

S. wangpeishanii 


 Selaginella  Homostachys   
Zhang &  stat. nov. S
elaginella
Homo
-
stachys  Lectotype (desig-
nated here): S
elaginella
pallidissima 
S
elaginella
Complanatae
Lectotype (designated
here): S
elaginella
denticulata 
= Selaginella

Repentes

 – Lecto-
type (designated here):
Selaginella helvetica

S
elagi-
nella
ciliaris  S. pallidissima 
S. Heterostachys
S.Homostachys
         
 
S. Homoeophyllae 
S.DichotropaeS. Heterophyl-
lumS. sStachygyn-
andrum


   

S. denticulata S. helvetica



Selaginella




Selaginella longistrobilina



Zhou & Zhang • Classification of Selaginella
TAX ON
–
Version of Record
 Selaginella Oligomacrosporangiatae
 
 Didiclis Oligomacrosporangiatae 
     
 Oligomacrosporangiata 

  

Selaginella fissidentoides


Selaginella

Monostelicae
 
 


Lectotype (designated here):
Selaginella fis-
sidentoides

.
= Selaginella

Erectae


Selaginella inaequali-
folia

= Selaginella

Scandentes

 
Selaginella

Scan-
dentes

– Lectotype
(designated here):
Selaginella laevigata

S. will-
denowii

= Selaginella
Sarmentosae
Lectotype (designated here):
Selaginella inaequalifolia

= Selaginella

Pleurophyllae

Lectotype (designated here):
Selaginella willde-
nowii

= Selaginella

Bisulcatae


Selaginella bisulcata

S
elaginella
 Suberosae 
S
elaginella
suberosa
S
elaginella
 Pleiostelicae
Lycopodioi-
des Pleiostelica
Pleiostele

S
elaginella
uncinata

Selaginella  Suberosae
S. suberosa 
S. Suberosae
Selaginella inaequalifolia
 

S.

Erectae

 
S.
 Sarmentosae 
Selaginella
willdenowii
 
S.

Pleurophyllae



S. inaequalifolia
S. 
Oligomacrosporangiatae
      
S
elaginella
Pleiostelicae 
S.

Pleiostelicae



S
elaginella
fissidentoides





 S. braunii  S. delicatula
S. pennataS. pervilleiS. siamensis
S. willdenowii
 

   Selaginella  Tetragonostachyae 
& 
LycopodiumTetragonostachya
  S
elaginella
Tetragonostachyae   
S.
Tetragonostachyae   
Tetragonos-
tachyaLycopodioides Tetragonostachya 

    Lycopodium tetragonostachyum 
S
elaginella
tetragonostachya 

Lycopodium Planifolia
S
elaginella
Planifoliae
    
Lectotype (designated here): Lycopodium tetrago-
nostachyum 

Selaginella

Pronae

– Lectotype (designated here):
Selaginella ciliaris Spring.
= Selaginella

Proniflorae


Selaginella proniflora

.
    
SelaginellaTetragonostachyae

  Selaginella

L. tetragonostachyum
  

S.

Proniflorae


Proniflorae

S. proniflora

S
elaginella
compta
   



Selaginella Stachygynandrum 
Stachygynandrum

Lycopodium  Stachygynandrum

Lycopodium  Stachygynandrum


TAX ON
–Zhou & Zhang • Classification of Selaginella
Version of Record
Lyco-
podium  Stachygynandrum 
StachygynandraDidi-
clis Stachygynandrum 
  

S
elaginella
flabellata
S
elaginella
Heterophyllum
Lectotype
(designated here): S
elaginella
flabellata
Stachygynandrum 
Lycopodium  L. denticulatum L. f la-
bellatum L. helveticum 
 L. flabellatum   


S
elaginella
 Stachygynandrum  
  S
elaginella
Articulatae
S
elaginella
 Rosulatae
S
elaginella
Suberosae 

 
     



    

 Selaginella  Ascendentes 
&  stat. nov.
Selaginella

Ascendentes
Lectotype (designated here):
S
elaginella
alopecuroides 
S
elaginella
Radicantes
Lectotype (designated here): S
elaginella
alopecuroides

Selaginella alopecuroides

S.

Ascendentes

S.

Radi-
cantes



 
     

elaginella
doederleinii S. frondosa   
 S. commutata 
 S. trichophyllaS. scabrifolia

Selaginella rolandi-principis   



 Selaginella  Austroamericanae 
& sect. nov.

Selaginella hartwegiana

= Selaginella

Redivivae
 
– Lectotype (designated
here):
Selaginella ciliata
 S
.
novae-
hollandiae
      


  Selaginella hartwegiana S. novae-
hollandiae


  
 

S. hartwegiana

 Selaginella Circinatae 
    stat. nov.Lycopodium
Circinatae     
CircinataS
elaginella
Circina-
tae
 Lecto type (designated here): S
elaginella
invol-
vens   

Selaginella

Rosulatae


S.

Rosulatae

 Lycopodioides Rosu-
latae 
S
elaginella
involvens     
= Selaginella

Caulescentes

 
S. 
Caulescentes
 

Selaginella caulescens
 S. involvens
 

Selaginella

Caulescentes
   
    


S. caulescens

S. asperula


S.

Circinatae

S.

Articulatae


Selaginella
Circinatae 

S.

Rosulatae


& 




S
elaginella
davidii S. gebaueri-
ana  S. moellendorffii 
S. involvens


 Selaginella Heterophyllae 
Lectotype (designated here):
Selaginella flexuosa


1137
Zhou & Zhang • Classification of Selaginella
TAX ON
–
Version of Record

S
elaginella
Enodes 
Lectotype (designated here): S
elaginella
flexuosa 

Selaginella

Adscendentes

 –
Lecto type (designated here):
Selaginella flexuosa


Selaginella

Persistentes

– Lectotype (designated
here):
Selaginella flexuosa


S
elaginella
Dichotropae  
Lectotype (des-
ignated here):
Selaginella flexuosa


Selaginella

Pleiomacrosporangiatae

  
Lecto type (designated here):
Selaginella flexuosa


Selaginella

Monostelicae

Lectotype
(designated here):
Selaginella flexuosa

S
elaginella
Continuae 
 
S.
Continuae 

Lectotype (designated here): S
elaginella
microphylla 
S
elaginella
Pusillae 
Lectotype (designated here):
S
elaginella
microphylla 

Selaginella

Resupinatae

 – Lectotype (designated
here):
Selaginella stenophylla

= Selaginella

Decumbentes

– Lectotype (designated here): S
elaginella
microphylla

S
elaginella
  Stachygynandrum
Lycopodioides 
Stachygynandrum
S
elaginella
flabel-
lata 
Selaginella
flexuosa
 
S. Heterophyllae

    Selaginella  
Resupina-
tae

S. stenophylla    

Diplostachyum tenellum 
Diplostachyum 

D. helveticum    
Selaginella tenella




      
Selaginella

Monostelicae

S.

Monostelicae
  


 Selaginella
Dichotropae

S.

Heterophyllae




 


 

   
nS. simplex 
nSelaginella erythropus
 S. flabellata
 Selaginella  Pallescentes
     S. pallescens 



Selaginella apoda 

  

S. apodan
S. pallescens  nS. pulcherrima
n  



 SelaginellaPlagiophyllae  
Zhang & stat. nov. S
elaginella
Pla-
giophyllaeLectotype (des-
ignated here): S
elaginella
biformis 
Selaginella biformis  
S. Plagiophyllae   

Selaginella
biformis   

  
     

S. biformis  
    
n
 Selaginella  Proceres  
& stat. nov.  Selginella Proceres
 Lec-
totype (designated here): S
elaginella
oaxacana 
       
S
elagi-
nella
martensii   
S. oaxacana


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TAX ON
–Zhou & Zhang • Classification of Selaginella
Version of Record
Alston, A.H.G., Jermy, A.C. & Rankin, J.M. Selagi-
nella Bull. Brit. Mus. (Nat. Hist.), Bot.

Arrigo, N., Therrien, J., Anderson, C.L., Windham, M.D., Haufler,
C.H. & Barker, M.S. 
 
SelaginellaTetragonostachys. Amer. J. Bot.

Baker, J.G. SelaginellaJ. Bot.

Baker, J.G. SelaginellaJ. Bot.

Baker, J.G. SelaginellaJ. Bot.

Baker, J.G. Handbook of the fern-allies: A synopsis of the genera
and species of the natural orders Equisetaceae, Lycopodiaceae,
Selaginellaceae, Rhizocarpeae

Banks, J.A.    Selaginella   
Annual Rev. Pl. Biol.

Banks, J.A., Nishiyama, T., Hasebe, M., Bowman, J.L., Gribskov,
M., dePamphilis, C., Albert, V.A., Aono, N., Aoyama, T.,
Ambrose, B.A., Ashton, N.W., A xtell, M.J., Barker, E., Barker,
M.S., Bennetzen, J.L., Bonawitz, N.D., Chapple, C., Cheng,
C., Correa, L.G., Dacre, M., DeBarry, J., Dreyer, I., Elias,
M., Engstrom, E.M., Estelle, M., Feng, L., Finet, C., Floyd,
S.K., Frommer, W.B., Fujita, T., Gramzow, L., Gutensohn,
M., Harholt, J., Hattori, M., Heyl, A., Hirai, T., Hiwatashi, Y.,
Ishik awa, M., Iwata, M., K arol, K.G., Koeh ler, B ., Kolukisaogl u,
U., Kubo, M., Kurata, T., Lalonde, S., Li, K., Li, Y., Litt, A.,
Lyons, E., Manning, G., Maruyama, T., Michael, T.P., Mikami,
K., Miyazaki, S., Morinaga, S., Murata, T., Mueller-Roeber, B.,
Nelson, D.R., Obara, M., Oguri, Y., Olmstead, R.G., Onodera,
N., Petersen, B.L., Pils, B., Prigge, M., Rensing, S.A., Riano-
Pachon, D.M., Roberts, A.W., Sato, Y., Scheller, H.V., Schulz,
B., Schulz, C., Shakirov, E.V., Shibagaki, N., Shinohara, N.,
Shippen, D.E., Sorensen, I., Sotooka, R., Sugimoto, N., Sugita,
M., Sumikawa, N., Tanurdzic, M., Theissen, G., Ulvskov, P.,
Wakazuki, S., Weng, J.K., Willats, W.W., Wipf, D., Wolf,
P.G., Yang, L., Zimmer, A.D., Zhu, Q., Mitros, T., Hellsten,
U., Loque, D., Otillar, R., Salamov, A., Schmutz, J., Shapiro,
H., Lindquist, E., Lucas, S., Rokhsar, D. & Grigoriev, I.V. 
Selaginella    
Science

Braun, A.       Appendix
plantarum novarum et minus cognitarum in Horto regio botanico
Berolinensi coluntur 1857

Chang, C.-Y., Liu, Q.-R. & Chen, X.-D. 
SelaginellaJ . Chin. Electron Microscopy
Soc.
Chang, C.-Y., Liu, Q.-R. & Chen, X.-D.     
Selaginella. J . Chin. Electron Microscopy
Soc.
Chang, H.-M., Chiou, W.-L. & Wang, J.-C. Flora of Taiwan:
Selaginellaceae  
 
Chu, W.M. Flora
Yunnanica
Da Silva Almeida, J.R .G., de Sá, P.G.S., de Oliveira Macedo, L.A.R .,
de Siqueira Filho, J.A., de Oliveira, V.R. & Filho, J.M.B. 
SelaginellaJ. Me d.
Pl. Res.
Dixit, R.D. Selaginellaceae of India

Giorgi, A.F.D., Holderegger, R. & Schneller, J.J.  
SelaginellaAmer.
Fern J.

Graustein, J.E. SelaginellaBot. Gaz.

Greville, R.K. & Hooker, W.J. Bot. Misc.

Gu, W., Song, J.-Y., Cao, Y., Sun, Q.-W., Yao, H., Wu, Q.-N., Chao,
J.-G., Zhou, J.-J., Xue, W.-D. & Duan, J.-A. 

PLoS ONE
   
Heidel, B. & Handley, J.  (L.) Beauv.
ex Mart. & Schrank (club spikemoss): A technical conservation
assessment


Hellwig, R.L. Selaginellae
Ann. Missouri Bot. Gard.

Hieronymus, G. & Sadebeck, R.      
Die natürlichen Pflanzenfamilien

Huang, T.C. Spore flora of Taiwan. 

Jermy, A.C. SelaginellaFern Gaz. 

Jermy, A.C.  
The families and genera of vascular plants
Pteridophytes and gymnosperms   

Jermy, A.C., Jones, K. & Colden, C. 
SelaginellaJ. Linn. Soc., Bot.

Jones, G.N. Selaginella douglasiiAmer.
Fern
J.
Korall, P. & Kenrick, P. 
rbcLAmer. J. Bot.
 
Korall, P. & Kenrick, P. 
    
 Molec. Phylo-
gen. Evol. 
Korall, P. & Taylor, W.A. 
   
 
Grana
ACKNOWLEDGMENTS
         
        
         
          

             
    
 
       
    


LITERATURE CITED

1139
Zhou & Zhang • Classification of Selaginella
TAX ON
–
Version of Record
Korall, P., Kenrick , P. & Therrien, J.P.   
   
rbcL Int. J. Pl. Sci. 
 
Kung, H.S. Flora Sichuanica

Kuntze, O. Revisio generarum plantarum 

Kunze, G. Linnaea
Lamarck, J.B. & Mirbel, C.F.B. de Histoire naturelle des végé-
taux, classés par familles         

Li, Q., Sun, X.-H. & Liu, J.-X. 
SelaginellaJ. Capital Norm. Univ. (Nat. Sci. Ed.)

Liu, B.-D., Bao, W.-M. & Aur, C.W.  
Bull. Bot.
Res., Harbin
Liu, J.-X. & Yan, X.-F.          
Chin. Bull. Bot.
Liu, J.-X. & Yan, X.-F.  
Chin. Bull.
Bot.
Liu, J.-X., Sun, X.-H., Dong, K., Zhang, J.-M., Li, Q. & Zhang X.-C.
   
 J. Chin. Electron Microscopy
Soc.
Liu, J.-X., Zhang, X.-C., Sun, X.-H. & Jiang, Y. 
J. Chin.
Electron Microscopy Soc.
Liu, J.-X., Sun, X.-H., Jin, Y., Lu, L., Yu, X.-X. & Zhang, X.-C.

  Acta Bot. Yunnan.
Liu, X.-R., Wang, G.-Z., Zhang, Q.-Y., Zhu, Y.-Q., Sun, X.-H.
& Zhao, Y.-Y.  
J. Jishou Univ. (Nat. Sci.)
Ludwig, C.G. & Böhmer, G.R. Definitiones generum plantarum


Marcon, A.B., Barros I.C.L. & Guerra, M. 

SelaginellaAnn. Bot. (Oxford)
 
McNeill, J., Barrie, F.R., Buck, W.R., Demoulin, V., Greuter, D.L.,
Hawksworth, D.L., Herendeen, P.S., Knapp, S., Marhold,
K., Prado, J., Proud’Homme van Reine, W.F., Smith, J.F.,
Wiersema, J.H. & Turland, N.J. (eds.) International Code
of Nomenclature for algae, fungi and plants (Melbourne Code):
Adopted by the Eighteenth International Botanical Congress, Mel-
bourne, Australia, July 2011. 

Mickel, J.T. & Hellwig, R.L. 
SelaginellaAmer. Fern J.
 
Mickel, J.T., Smith, A.R. & Valdespino, I.A. Selaginella
The pteridophytes
of Mexico

Minak i, M. Selaginella
Pollen & Spores 
Moore, T. Index filicum

Palisot-Beauvois, A.M.F.J. Mag. Encycl.

Pfeiffer, L.G.K. Nomenclator botanicus

 
Punt, W., Hoen, P.P., Blackmore, S., Nilsson, S. & Le Thomas, A.
Rev. Palaeobot.
Palynol.   
Reeve, R.M. Selaginella
Rhodora
Reichenbach, H.G.L.  Conspectus regni vegetabilis 

Rothmaler, W. Feddes Reper t. Spec. Nov.
Regni Veg.
Schulz, C., Homberg, J. & Stützel, T.  
SelaginellaEricetorumSyst. Bot. 

Setyawan, A.D. Selaginella
Nusantara Biosci.

Singh, S.K., Yadav, B.B., Srivastava, M., Shukla, P.K. & Srivastava,
G.K. Selaginella
Grana
 
Soják, J. G
Preslia
Sprengel, K.P.J.  Anleitung zur Kenntniss der Gewächse

Spring, A.F. Flora

Spring, A.F. 
   Flora brasiliensis

Spring, A.F. Mém.
Acad. Roy. Sci. Belgique
Stefanović, S., Rakotondrainibe, F. & Badré, F. 
Flore de Madagas-
car et des Comores

Sun, Q.-W., Zhao, J.-H. & Zhang, L.-B.  Selaginella daozhe-
nensis 
Phytotaxa 

Sun, X.-H., Zhang, J.-M., Jiang, Y., Liu, J.-X. & Zhang, X.-C. 

SelaginellaJ. Capital Norm. Univ.
(Nat. Sci. Ed.)
Tagawa, M. & Iwatsuki, K.     
Flora of Thailand  

Takamiya, M. 
SelaginellaJ. Pl. Res.
 
Thomas, B.A. Rev.
Palaeobot. Palynol.

Tryon, A.F.     Selaginella 
Ann. Missouri Bot. Gard.

Tryon, A.F. & Lugardon, B. Spores of the Pteridophyta: Surface,
wall structure, and diversit y based on electron microscope studies

 
Tryon, R.M. Selaginella rupestris Ann. Missouri
Bot. Gard.
Tryon, R.M. & Tryon A.F. Ferns and allied plants: With special
reference to tropical America.
 
Tschermak-Woess, E. & Doležal-Janish, R. 

Cyrtomium falcatumOesterr. Bot. Z.


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–Zhou & Zhang • Classification of Selaginella
Version of Record
Tzvelev, N. Selaginella
Novosti Sist. Vyssh. Rast. 
Valdespino, I.A. 
Flora of North America
Pteridophytes and gymnosperms

Valdespino, I.A. Selaginella
Brittonia

Walton, J. & Alston, A.H.G. 
Manual of pteridology
 
Warburg, O. Monsunia

Weakley, A.S. Flora of the southern and mid-Atlantic States

Xia, Y., Dai, X.-L., Yan, Y.-H., Cao, J.-G. & Wang, Q.-X. 
  
Acta Bot. Boreal. -Occid. Sin.
Zhang, L.-B., Wang, P.-S. & Wang, X.-Y. Selaginella longistro-
bilina
Selaginella prostrata       
Novon 
Zhang, L.-B., Sun, Q.-W. & He, H.  Selaginella wangpeisha-
nii 
Phytotaxa

Appendix 1.
Selaginella
A–L,

M–X,

A, M,
S.Boreoselaginella; S. sanguinolentaChu 23322
B, C, N, O,
S.  Tetragonostachys
B, N,
S indica S. 
Homoeophyllae
Chu & Wu 2598 
C, O.
S. remotifolia
 S. 
Articulatae

Gao & al. DJY2931

D, P,
S.PulviniellaS. stauntoniana 
Zhao
169 
E–J, Q–V,
S.Heterostachys.
E, Q,
S. heterostachys S. HeterostachysZhou & Jü DJY4117  
F,R,
S. ciliaris  S. Tetragonostachyae
Jiang 310

G, S,
S. pallidissima S. Homostachys
Qinghai-Xizang Exped. 13303
H, T,
S. braunii S. 
Oligomacrosporangiatae

Zhou & Jiang
217

I, U,
S. pseudopaleifera S. 
Oligomacrosporangiatae

Zhang
& al.
6341

J, V,
S. pen-
nata  S. 
Oligomacrosporangiatae

He & Jiang 303

K, L, W, X.
S. Stachygynandrum
K, W.
S. involvens  S. Circinatae
Zhou & al. DJY03558

L, X,
S. doederleinii S. 
Ascendentes


Jü & al. HGX12933

Zhang, X.-C., Kato, M. & Nooteboom, H.P. 
Flora
of China  

Zhao, Y.-Y., Liu, X.-R., Zhu, Y.-Q., Sun, X.-H. & Liu, J.-X. 
  Selaginella
J. Chin. Electron Microscopy Soc
Zhao, Y.-Y., Zhang, Q.-Y., Jin, Y., Sun, X.-H. & Liu, J.-X. 

Biotechnol. Bull.
Zhou, X.-M., Zhang, Y.-J., Li, Z.-L., He, Z.-R. & Liu, Y.-J. 
      Selaginella 
J. Chin. Electron Microscopy Soc.
Zhou, X.-M., Rothfels, C.J., Zhang, L., He, Z.-R., Le Péchon, T.,
He, H., Lu, N.T., Knapp, R., Lorence, D., He, X.-J., Gao, X.-F.
& Zhang, L.-B. 
 Selaginella  
Cladistics

Zhou, X.-M., Jiang, L.-J., Zhang, L., Gao, X.-F., He, Z.-R. & Zhang,
L.-B.  Selaginella
     Phytotaxa  
   
Zhu, Y.-Q., Wang, G.-Z., Liu, X.-R. Zhang, Q.-Y., Sun, X.-H. & Liu,
J.-X. 
J. Jishou Univ. (Nat. Sci.)

Vol. 64 (6) • December 2015
International Journal of Taxonomy, Phylogeny and Evolution
Electronic Supplement to
A classification of Selaginella (Selaginellaceae)
based on molecular (chloroplast and nuclear),
macromorphological, and spore features
Xin-Mao Zhou & Li-Bing Zhang
Taxon 64: 1117–1140

S1
Electr. Suppl. to: Zhou & Zhang • Classif ication of Selaginella
TAX ON
64 (6) • December 2015
Fig. S1.
Maximum likelihood phylogeny of Selaginella based on plastid rbcL and nuclear ITS sequences of 394 accessions representing ca. 200
species. Geographical provenances of newly included samples of Selaginella are indicated in red and those f rom Gen Bank are indicated in other
colors. Micrographs show mor phology of megaspores and microspores. Outgroups are removed. Support values are not shown (see Fig. 6).
0.05
Subg. Stachygynandrum Subg. Heterostachys
Subg. Pulviniella
Subg. Selaginella
Subg. Ericetorum
Subg. Boreoselaginella
Sect. Lyallia
Sect. Myosurus
Sect. Megalosporarum
Sect. Articulatae
Sect.
Homoeophyllae
Sect. Lepidophyllae
Sect. Oligomacrosporangiatae
Sect. Auriculate
Sect. Homostachys
Sect. Tetragonostachyae
Sect. Heterostachys
Sect. Ascendentes
Sect. Austroamericanae
Sect. Heterophyllae
Sect. Pallescentes
Sect. Poceres
Sect. Circinatae
Sect. Plagiophyllae
Hainan, China
Hainan, China
Hainan, China
Introduced to Costa Rica
Cult.
Beijing, China
Venezuela
Bac Kan, Vietnam
Bac Kan,Vietnam
Bac Kan, Vietnam
Ha Giang, Vietnam
Okinawa, Japan
Kagoshima, Japan
Sichuan, China
Yunnan, China
Phu Tho, Vietnam
Cult.
Yunnan, China
Sichuan, China
Sichuan, China
Sichuan, China
Guizhou, China
Yunnan, China
Guizhou, China
Yunnan, China
Yunnan, China
Guizhou, China
Heredia, Costa Rica
Oaxaca, Mexico
Oaxaca, Mexico
Cult.
Oaxaca, Mexico
San Luis Potosi, Mexico
Unknown
Virginia, USA
Tennessee, USA
Unknow
San Luis PotosÌ, Mexico
Cult.
Hidalgo, Mexico
Cult.
Unknown
Unknown
San Jose, Costa Rica
Guanacaste, Bagaces
Hidalgo, Mexico
Guanacaste, Costa Rica
Jalisco, Mexico
Heredia, Costa Rica
San Luis PotosÌ, Mexico
Loja, Ecuador
Ecuador
Zamora-Chinchipe, Ecuador
Guanacaste, Costa Rica
Heredia, Costa Rica
Carchi, Ecuador
Unknown
Cult.
Heredia, Costa Rica
Unknown
French guiana
Guanacaste, Costa Rica
Heredia, Costa Rica
Lesser Antilles, Grenada
Heredia, Costa Rica
Unknown
Unknown
Unknown
San José, Costa Rica
Borneo
Peninsular Malaysia
Unknown
Unknown
Cult.
Hainan, China
Kagoshima, Japan
Yunnan, China
Guangxi, China
Hainan, China
Yunnan, China
Unknown
Yunnan, China
Yunnan, China
Peninsular Malaysia
Bomeo
Unknown
San José, Costa Rica
*
*
*
*
Cult.
*
*
*
*
*
Guangxi, China
Guizhou, China
Sichuan, China
Sichuan, China
Sichuan, China
Sichuan, China
Sichuan, China
Sichuan, China
Sichuan, China
Sichuan, China
Guizhou, China
Hoa Binh, Vietnam
Ha Giang, Vietnam
Yunnan, China
Guizhou, China
Guangxi, China
Guangxi, China
Yunnan, China
Yunnan, China
Yunnan, China
Sichuan, China
Sichuan, China
Sichuan, China
Guangxi, China
Guizhou, China
Guangxi, China
Yunnan, China
Yunnan, China
Yunnan, China
Tokyo, Japan
Lang Son, Vietnam
Lang Son, Vietnam
Hainan, China
Hainan, China
Guangxi, China
Thai Nguyen,Vietnam
Lang Son, Vietnam
Lang Son,Vietnam
Guangxi, China
Guangxi, China
Mie, Japan
Sichuan, China
Guangxi, China
Hoa Binh, Vietnam
Phu Tho, Vietnam
Ha Giang, Vietnam
Yunnan, China
Sichuan, China
Sichuan, China
Guizhou, China
Guangxi, China
Yunnan, China
Bac Kan,Vietnam
Yunnan, China
Yunnan, China
Yunnan, China
Cao Bang, Vietnam
Hainan, China
Guangxi, China
Sichuan, China
Sichuan, China
Bac Kan, Vietnam
Bac Kan, Vietnam
Vinh Phuc, Vietnam
Yunnan, China
Yunnan, China
Guangxi, China
Guangdong, China
Guangxi, China
Guizhou, China
Guizhou,China
Guizhou,China
Guizhou, China
Huahine, Society Is.
Caroline Is., Kosrae, F.S.M.
Caroline Is., Kosrae, F.S.M.
Maui, Hawaiian Is.
Kauai, Hawaiian Is.
Caroline Is., Pohnpei, F.S.M.
Caroline Is., Pohnpei, F.S.M.
Yunnan, China
Yunnan, China
Yunnan, China
Yunnan, China
Guangxi, China
Okinawa, Japan
Hainan, China
Peninsular Malaysia
Guangdong, China
Bac Kan, Vietnam
Sichuan, China
Sichuan, China
Guizhou, China
Guizhou, China
Mie, Japan
Sichuan, China
Sichuan, China
Yunnan, China
Sichuan, China
Yunnan, China
Yunnan, China
Yunnan, China
Sichuan, China
Akita, Japan
Unknown
Taiwan, China
Taiwan, China
Taiwan, China
Taiwan, China
Taiwan, China
Taiwan, China
Taiwan, China
Taiwan, China
Taiwan, China
Sichuan, China
Sichuan, China
Sichuan, China
Sichuan, China
Taiwan, China
Tokyo, Japan
Taiwan, China
Taiwan, China
Oregon, USA
Yunnan, China
Yunnan, China
Yunnan, China
Sichuan, China
Cult. in Thailand
Yunnan, China
Hainan, China
Hubei, China
Hoa Binh, Vietnam
Kagoshima, Japan
Yunnan, China
Ha Giang, Vietnam
Yunnan, China
Cult.
Guizhou, China
Guangxi, China
Peninsula Malaysia
Puntarenas, Costa Rica
Cult.
Peninsula Malaysia
Cult.
Puntarenas, Costa Rica
Puntarenas, Costa Rica
Bac Kan, Vietnam
Sichuan, China
Sichuan, China
Phu Tho, Vietnam
Yunnan, China
Yunnan, China
Yunnan, China
Unknown
Sichuan, China
Tokyo, Japan
Sichuan, China
Unknown
Fianarantsoa, Madagascar
Yunnan, China
Yunnan, China
Yunnan, China
Yunnan, China
Yunnan, China
Yunnan, China
Yunnan, China
Dhofar, Oman
Unknown
Unknown
d'Andohahela, Madagascar
d'Andohahela, Madagascar
Unknown
Henan, China
Beijing, China
Okinawa, Japan
Unknown
Unknown
Sichuan, China
Xizang, China
Texas , USA.
Unknown
Isiolo District, Kenya
Matale District., Sri-Lanka
Jalisco, Mexico
California, USA
California, USA
Arizona, USA
Utah, USA
Idnho, USA
British Columbia, Canada
Washington, USA
British Columbia, Canada
North Carolina, USA
North Carolina, USA
Ontario, Canada
Hokkaido, Japan
Alaska, USA
Utah to Arizona, USA
Illinois , USA
Texas, USA
Hidalgo, Mexico
Texas, USA
Oaxaca, Mexico
Isiolo District, Kenya
Fianarantsoa, Madagascar
Tulear, Madagascar
Unknown
San Luis Potosi, Mexico
Carchi, Ecuador
Florída, Bolivia
Unknown
Boyocá Dept., Colombia
California, USA
California, USA
Louisiana, USA
Jalisco, Mexico
Colorado, USA
Washington, USA
Tamegdoult, Morocco
South carolina, USA
North Carolina, USA
Pretoria, South Africa.
Hassan District, India
Cult.
Yunnan, China
Sichuan, China
Yunnan, China
Yunnan, China
Yunnan, China
Quezaltenango, Guatemala
California, USA
California, USA
Oregon, USA
British Columbia, Canada
Oregon, USA
California, USA
California, USA
New Mexico, USA
Yunnan, China
Nagano, Japan
Nagano, Japan
Colorado, USA
Idaho, USA
Washington, USA
Idaho, USA
Texas, USA
Moreles, Mexico
Texas, USA
Arizona, USA
Unknown
Arizona, USA
California, USA
California, USA
California, USA
Texas, USA
New Mexico, USA
New Mexico, USA
San Luis PotosÌ, Mexico
New Mexico, USA
Unknown
Colorado, USA
Kwale District, Kenya
California, USA
California, USA
Arizona, USA
Colorado, USA
Cult.
Cult.
Cult.
Sichuan, China
Shizuoka, Japan
Yunnan, China
Unknown
Unknown
Unknown
Pichincha, Ecuador
Unknown
Unknown
Heredia, Costa Rica
Jalisco, Mexico
Unknown
Heredia, Costa Rica
Heredia, Costa Rica
Heredia, Costa Rica
Puntarenas, Costa Rica
Puntarenas, Costa Rica
Cult.
Unknown
Unknown
Unknown
Fianarantsoa, Madagascar
Fianarantsoa, Madagascar
d'Antsiranana, Madagascar
Tasmania, Australia
Tasmania, Australia
Hawaii
Japan
Ontario, Canada
Unknown
Sichuan, China
Sichuan, China
Sichuan, China
Yunnan, China
Xizang, China
Unknown
Unknown
S. biformis
S. biformis
S. biformis
S. biformis
S. biformis
S. biformis
S. biformis
S. biformis
S. involvens
S. involvens
S. involvens
S. involvens
S. involvens
S. moellendorffii
S. moellendorffii
S. moellendorffii
S. moellendorffii
S. moellendorffii
S. davidii
S. gebaueriana
S. gebaueriana
S. gebaueriana
S. gebaueriana
S. gebaueriana
S. novae-hollandiae
S. oaxacana
S. oaxacana
S. oaxacana
S. martensii
S. reflexa
S. apoda
S. apoda
S. apoda
S. acanthostachys
S. harrisii
S. pallescens
S. harrisii
S. pulcherrima
S. pallescens
S. pallescens
S. pallescens
S. pallescens
S. pallescens
S. pallescens
S. pallescens
S. pallescens
S. nothohybrida
S. hartwegianum
S. novae hollandiae
S. flexuosa
S. anceps
S. anceps
S. microphylla
S. haematodes
S. erythropus
S. bombycina
S. bombycina
S. radiata
S. attirense cf.
S. umbrosa
S. umbrosa
S. flabellata
S. flagellata
S. flagellata
S. simplex
S. moritziana
S. attirense cf.
S. porphyrospora
S. alopecuroides
S. intermedia
S. firmuloides
S. longipinna
S. kerstingii
S. doederleinii
S. doederleinii
S. doederleinii
S. commutata
S. scabrifolia
S. trachyphylla
S. frondosa
S. frondosa
S. superba
S. roxburghii
S. brooksii
S. longiaristata
S. gebaueriana
S. martensii
S. involvens
S. drepanophylla
S. drepanophylla
S. chrysocaulos
S. labordei
S. labordei
S. labordei
S. labordei
S. sichuanica
S. sichuanica
S. sichuanica cf.
S. daozhenensis
S. megaphylla
S. megaphylla
S. megaphylla
S. megaphylla
S. megaphylla
S. megaphylla
S. chaetoloma
S. chaetoloma
S. chaetoloma
S. bodinieri
S. bodinieri
S. bodinieri
S. bodinieri
S. bodinieri
S. bodinieri
S. xichouensis
S. kurzii
S. kurzii
S. boninensis
S. boninensis
S. boninensis
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. heterostachys
S. sp.
S. sp.
S. leptophylla
S. leptophylla
S. leptophylla
S. leptophylla
S. leptophylla
S. monospora cf.
S. tenuifolia
S. effusa var. dulongjiangensis
S. trichophylla
S. trichophylla
S. trichophylla
S. trichophylla
S. amblyphylla cf.
S. amblyphylla
S. amblyphylla
S. decipiens
S. decipiens
S. monospora
S. monospora
S. monospora
S. effusa
S. effusa
S. effusa
S. effusa
S. sp.
S. sp.
S. sp.
S. sp.
S. sp.
S. sp.
S. arbuscula
S. arbuscula
S. kanehirae
S. kanehirae
S. repanda
S. repanda
S. repanda
S. repanda
S. albociliata
S. lutchuensis
S. ciliaris
S. xipholepis
S. xipholepis
S. compta
S. compta
S. compta
S. compta
S. compta
S. ciliaris
S. tama-montana
S. prostrata
S. prostrata
S. pallidissima
S. laxistrobila
S. laxistrobila
S. laxistrobila
S. helvetica
S. helvetica
S. denticulata
S. tama-montana
S. pseudonipponica
S. pseudonipponica
S. pseudonipponica
S. pseudonipponica
S. pseudonipponica
S. pseudonipponica
S. sp
S. helvetica
S. jugorum
S. nipponica
S. nipponica
S. nipponica
S. nipponica
S. nipponica
S. matsuensis
S. nipponica
S. pseudonipponica
S. pseudonipponica
S. douglasii
S. mairei
S. mairei
S. mairei
S. mairei
S. braunii
S. braunii
S. braunii
S. braunii
S. pseudopaleifera
S. limbata
S. helferi
S. helferi
S. helferi
S. willdenowii
S. willdenowii
S. willdenowii
S. uncinata
S. uncinata cf.
S. uncinata
S. uncinata
S. douglasii
S. plana
S. plana
S. hoffmannii
S. hoffmannii
S. wallichii
S. delicatula
S. delicatula
S. mayeri
S. picta
S. picta
S. picta
S. siamensis
S. pervillei
S. pennata
S. pennata
S. bisulcata
S. bisulcata
S. bisulcata
S. bisulcata
S. bisulcata
S. imbricata
S. imbricata
S. pilifera
S. helioclada
S. stauntoniana
S. stauntoniana
S. stauntoniana
S. tamariscina
S. tamariscina
S. tamariscina
S. pulvinata
S. pulvinata
S. lepidophylla
S. digitata
S. lepidophylla
S. phillipsiana
S. wightii
S. landii
S. rupestris
S. asprella
S. leucobryoides
S. leucobryoides
S. utahensis
S. watsonii
S. densa
S. scopulorum
S. scopulorum
S. acanthonota
S. rupestris
S. rupestris
S. sibirica
S. sibirica
S. watsonii
S. wrightii
S. arsenei
S. sheldonii
S. peruviana
S. mutica
S. dregei
S. echinata
S. grisea
S. nivea
S. arsenei
S. sellowii
S. sellowii
S. sellowii
S. sartorii
S. cinerascens
S. cinerascens
S. arenicola
S. extensa
S. weatherbiana
S. oregana
S. balansae
S. tortipila
S. tortipila
S. caffrorum
S. indica
S. indica
S. indica
S. vardei
S. vardei
S. vardei
S. vardei
S. steyermarkii
S. hansenii
S. wallacei
S. wallacei
S. wallacei
S. wallacei
S. bigelovii
S. bigelovii
S. bigelovii
S. bigelovii
S. mutica var. limitanea
S. mutica var. limitanea
S. × neomexicana
S. × neomexicana
S. sartorii
S. rupincola
S. rupincola
S. rupincola
S. underwoodii
S. njamnjamensis
S. vardei
S. shakotanensis
S. wallacei
S. wallacei
S. wallacei
S. shakotanensis
S. densa
S. peruviana
S. landii
S. wrightii
S. arizonica
S. arizonica
S. arizonica
S. eremophila
S. eremophila
S. eremophila
S. kraussiana
S. kraussiana
S. kraussiana
S. remotifolia
S. remotifolia
S. remotifolia
S. remotifolia
S. fragilis
S. articulata
S. sericea
S. sericea
S. suavis
S. sulcata
S. eurynota
S. sertata
S. lingulata
S. arthritica
S. arthritica
S. silvestris
S. kunzeana
S. diffusa
S. diffusa
S. exaltata
S. myosurus
S. pygmaea
S. lyallii
S. moratii
S. polymorpha
S. gracillima
S. uliginosa
S. deflexa
S. selaginoides
S. selaginoides
S. selaginoides
S. sanguinolenta
S. sanguinolenta
S. sanguinolenta
S. sanguinolenta
S. sanguinolenta
S. nummularifolia