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A molecular phylogeny and new subgeneric classification of Sporobolus (Poaceae: Chloridoideae: Sporobolinae)

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The grass subtribe Sporobolinae contains six genera: Calamovilfa (5 spp. endemic to North America), Crypsis (10 spp. endemic to Asia and Africa), Psilolemma (1 sp. endemic to Africa), Spartina (17 spp. centered in North America), Sporobolus (186 spp. distributed worldwide), and Thellungia (1 sp. endemic to Australia). Most species in this subtribe have spikelets with a single floret, 1-veined (occasionally 3 or more) lemmas, a ciliate membrane or line of hairs for a ligule, and fruits with free pericarps (modified caryopses). Phylogenetic analyses were conducted on 177 species (281 samples), of which 145 species were in the Sporobolinae, using sequence data from four plastid regions (rpl32-trnL spacer, ndhA intron, rps16-trnK spacer, rps16 intron) and the nuclear ribosomal internal transcribed spacer regions (ITS) to infer evolutionary relationships and provide an evolutionary framework on which to revise the classification. The phylogenetic analysis provides weak to moderate support for a paraphyletic Sporobolus that includes Calamovilfa, Crypsis, Spartina, and Thellungia. In the combined plastid tree, Psilolemma jaegeri is sister to a trichotomy that includes an unsupported Urochondra-Zoysia clade (subtr. Zoysiinae), a strongly supported Sporobolus somalensis lineage, and a weakly supported Sporobolus s.l. lineage. In the ITS tree the Zoysiinae is sister to a highly supported Sporobolinae in which a Psilolemma jaegeri–Sporobolus somalensis clade is sister to the remaining species of Sporobolus s.l. Within Sporobolus s.l. the nuclear and plastid analyses identify the same 16 major clades of which 11 are strongly supported in the ITS tree and 12 are strongly supported in the combined plastid tree. The positions of three of these clades representing proposed sections Crypsis, Fimbriatae, and Triachyrum are discordant in the nuclear and plastid trees, indicating their origins may involve hybridization. Seven species fall outside the major clades in both trees, and the placement of ten species of Sporobolus are discordant in the nuclear and plastid trees. We propose incorporating Calamovilfa, Crypsis, Spartina, Thellungia, and Eragrostis megalosperma within Sporobolus, and make the requisite 35 new combinations or new names. The molecular results support the recognition of 11 sections and 11 subsections within Sporobolus s.l.; four sections are new: Airoides, Clandestini, Cryptandri, and Pyramidati; three sections are new combinations: Calamovilfa, Crypsis, and Spartina; four subsections are new combinations: Calamovilfa, Crypsis, Ponceletia, and Spartina; seven subsections are new: Actinocladi, Alterniflori, Floridani, Helvoli, Pyramidati, Spicati, and Subulati; 30 new combinations in Sporobolus: S. aculeatus, S. advenus, S. alopecuroides, S. alterniflorus, S. angelicus, S. arcuatus, S. bakeri, S. borszczowii subsp. acuminatus, S. borszczowii subsp. ambiguus, S. brevipilis, S. coarctatus, S. cynosuroides, S. densiflorus, S. factorovskyi, S. foliosus, S. hadjikyriakou, S. ×longispinus, S. maritimus, S. megalospermus, S. michauxianus, S. minuartioides, S. niliacus, S. pumilus, S. rigidus, S. rigidus var. magnus, S. spartinus, S. schoenoides, S. ×townsendii, S. turkestanicus, and S. vericolor; and five new names in Sporobolus: S. arenicola, S. ×eatonianus, S. hookerianus, S. mobberleyanus, and S. vaseyi are made. Lectotypes are designated for Crypsis factorovskyi, Heleochloa ambigua, and Torgesia minuartioides.
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

Sporobolus: phylogeny and classification
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

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INTRODUCTION
  

Urochondra
      

Zoysia 


A molecular phylogeny and new subgeneric classification of
Sporobolus (Poaceae: Chloridoideae: Sporobolinae)
Paul M. Peterson,1 Konstantin Romaschenko,1,2 Yolanda Herrera Arrieta3 & Jeffery M. Saarela4
 
 
 
 
 Paul M. Peterson, peterson@si.edu

DOI

Abstract
Crypsis
Psilolemma Spartina Sporobolus
Thellungia
                

              

Sporobolus 
 Crypsis Spartina  Thellungia   Psilolemma jaegeri   
Sporobolus somalensis 
Sporobolus      
Psilolemma jaegeri–Sporobolus somalensis   Sporobolus Sporobolus 

Crypsis
Triachyrum             
Sporobolus    
 Crypsis SpartinaThellungiaEragrostis megalosperma Sporobolus 

Sporobolus  ClandestiniCryptandriPyramidati    
 CrypsisSpartina Crypsis PonceletiaSpartina
  Pyramidati Spicati Subulati     
SporobolusS. aculeatus S. alopecuroides S. alterniflorus S. angelicus S. arcuatus   
acuminatus    ambiguus S. coarctatus S. cynosuroides S. densiflorus      S. folio
sus S. S. ×longispinus S. maritimus S. megalospermus S.minuartioides S. niliacus S. pumi
lus S. rigidus S. rigidusmagnusS. spartinus S. schoenoides
SporobolusS. arenicola S. ×eatonianus S. mobberleyanus 
Heleochloa ambigua Torgesia minuartioides.
Keywords
 CrypsisPogononeura Psilolemma
Spartina SporobolusThellungia
Supplementary Material




Sporobolus: phylogeny and classification
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  


  

Crypsis
Pogononeura
Spartina 
Sporobolus
 



 
  



Spartina


   Spartina 

   


Zoysia



Urochondra
  Zoysia




Crypsis UrochondraSporobolus

Sporobolus



 Sporobolus


 
Sporobolus
 

ChaetorhachiaEusporobolus
 Spo
robolusSporobolus   

Sporobolus
 
Sporobolus indicus

Sporobolus


Sporobolus floridanus


S. aeneus
Sporobolus 
  






         
   

    
Sporobolus   Calamo
 Crypsis Pogononeura Psilolemma
Urochondra  SpartinaZoysia
 




Spartina pectinata Spo
robolus airoides S. consimilus S. hetero
lepis S. junceusS. spicatus
       
 
Spartina alterniflora
S. maritima 
Crypsis alopecuroidesSpar
tina alternifloraS. anglicaS. densi flora
Sporobolus africanusS. fertilis
   S. indicus  S. pyramidalis 
 


 Sporobolus
  
Sporobolus

 
Sporobolus
Crypsis 
Sporobolus
Crypsis PogononeuraSpartina


  Sporobolus   





TAX ON


Sporobolus: phylogeny and classification
Version of Record (identical to print version).

 
Sporobolus


Sporobolus

   


Sporobolus   
 

MATERIALS AND METHODS
Taxon sampling. —

          






Sporobolus
 
Sporobolus


Sporobolus

Sporobolus



 Capeochloa
cintasericeaDan
thonia compressaRytidosperma penicellatum 
  
sophilaChasmanthium
latifolium





DNA extraction, amplification, and sequencing. —

 



 



 
Phylogenetic analyses. —
    

  
 
  



 

 
  
  






     

 






 

RES ULTS
Phylogenetic analyses. —
  
 

    
   
    Sporobolus aldabrensis
S. conrathii S. elongatus
S. floridanusS.maderaspatanus S. micranthus
S. mildbraedii
Zoysia pacifica
S. smutsiiS. tenacissimus



    






Analysis of ITS sequences. —
   
      


Sporobolus: phylogeny and classification
TAX ON

Version of Record (identical to print version).




ZoysiaUrochondra setu
losa.Sporobolus
Psilolemma jaegeri 
Sporobolus somalensis 
 
Sporobolus 
Crypsis SpartinaThellungia
Sporobolus    



  

 


Spartina alterniflora  S. anglica S. foliosa  S. mari
tima    
 S. ×caespitosa  S. ciliata  S. cyno
suroides  S. densiflora S. gracilis S. 
sis     S. patens S. pectinata 
S. spartinae 
Sporobolus
Sporobolus acinifolius –S. albicans –S. tenellus


Sporobolus
sis –S. robustus
S. consimilis 
 


S. palmeri

Sporobolus rigens 


Analysis of combined plastid sequences. —


 
Sporobolus  Crypsis Spartina
 ThellungiaPogononeura biflora 

Psilolemma

Urochon
dra–Zoysia
Sporobolus somalensis
Sporobolus
Sporobolus acinifolius–S. albicans–S. tenellus 


Table 1.
rps16 ,

  rps16  

 
 937 1236 964 954 4091 798
      
  
198

199

781
 
     
 6 6 6 6 6
      
 




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

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
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












 



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
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





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      
     
      

TAX ON


Sporobolus: phylogeny and classification
Version of Record (identical to print version).
0.1
Gymnopogon grandiflorus
Leptothrium rigidum
Eleusine indica
Chloris barbata
Cynodon plectostachyus
Melanocenchris abyssinica
Eragrostiella leioptera
Oropetium capense
Tripogon multiflorus
Urochondra setulosa
Zoysia japonica
Zoysia macrantha
Zoysia macrantha subsp. walshii
Psilolemma jaegeri Peterson 24247
Psilolemma jaegeri Peterson 24249
Sporobolus somalensis Boalev 317
Sporobolus somalensis Herruming 2022
Sporobolus farinosus Sporobolus pectinellus Fay 7131
Sporobolus festivus
Sporobolus infirmus
Sporobolus stapfianus Laegaard 15939
Sporobolus africanus Peterson 24024
Sporobolus africanus Peterson 24121
Sporobolus pyramidalis Peterson 24150
Sporobolus pyramidalis Senaratne 6082
Sporobolus pectinellus Peterson 23978
Sporobolus molleri Gereau 5790
Sporobolus molleri Rwaburindore 2183
Sporobolus fertilis Raulerson 775
Sporobolus fertilis Gould 13535
Sporobolus indicus Peterson 22025
Sporobolus jacquemontii Estrada 18964
Sporobolus jacquemontii Peterson 15902
Sporobolus berteroanus
Sporobolus indicus Peterson 7337
Sporobolus pseudairoides
Sporobolus myrianthus
Sporobolus creber
Sporobolus laxus
Sporobolus natalensis
Sporobolus sessilis
Sporobolus tenuissimus
Sporobolus blakei
Sporobolus diandrus
Sporobolus atrovirens Peterson 24661
Sporobolus atrovirens Peterson 22342
Sporobolus trichodes Rzedowski 39901
Sporobolus atrovirens Peterson 24729
Sporobolus atrovirens Rosalen 3381
Sporobolus albicans
Sporobolus tenellus
Sporobolus acinifolius
Sporobolus fimbriatus Peterson 24206
Sporobolus greenwayi
Sporobolus fimbriatus Peterson 24280
Sporobolus diffusus
Sporobolus brockmanii
Sporobolus fimbriatus Peterson 24241
Sporobolus confinis
Sporobolus phyllotrichus
Sporobolus pellucidus
Sporobolus nervosus
Sporobolus smutsii
Sporobolus montanus
Sporobolus ruspolianus McKinnon s.n.
Sporobolus ruspolianus Bally 15581
Sporobolus ruspolianus Flemming 2136
Crypsis schoenoides
Crypsis aculeata Soreng 5469
Crypsis aculeata Soreng7940
Crypsis alopecuroides
Sporobolus helvolus Laegaard 17063
Sporobolus helvolus Peterson 24217
Sporobolus mitchellii
Sporobolus humilis
Thellungia advena Belson 1930
Thellungia advena Lazarides 4185
Sporobolus consimilis Collenette 17
Sporobolus consimilis Peterson 24252
Sporobolus oxylepsis Schlieben 6158
Sporobolus robustus Laegaard 17398
Sporobolus sanguineus Bidgood 2397
Sporobolus sanguineus Gereau 6014
Sporobolus subglobosus
Sporobolus panicoides
Sporobolus dinklagei
Sporobolus purpurascens Peterson 9453
Sporobolus aeneus
Sporobolus acuminatus Guala 1372
Sporobolus acuminatus Irwin 11586
Sporobolus linearifolius
Sporobolus apicaulis
Sporobolus eylesii
Sporobolus stolzii Peterson 23946
Sporobolus stolzii Richards 21377
Sporobolus stolzii Peterson 24133
Sporobolus pilifer
Sporobolus bogotensis
Sporobolus lasiophyllus Peterson 21820
Sporobolus lasiophyllus Peterson 21879
Sporobolus junceus Strong 2332
Sporobolus junceus Thierat 25181
Sporobolus purpurascens Swallen 10179
Sporobolus purpurascens Trovart 25
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.81
0.90
0.95
1.00
1.00 1.00
0.90
1.00
1.00
1.00
1.00
1.00
0.94
1.00
1.00
0.89
0.82
1.00
1.00
1.00
0.89
0.85
1.00
0.85
0.99
1.00
0.99
1.00
0.68
0.95
0.57
1.00
1.00
0.99
0.99
1.00
0.97
0.92
1.00
1.00
0.80
1.00
1.00
1.00
1.00
0.97
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.98
1.00
0.67
1.00
1.00
1.00
0.98
1.00
0.96
0.97
1.00
0.95
1.00
99
95
78
83
96
54
77
100
97
100
68
100
100
100
69
100
65
100
100
71
82
100
98
1.00
75
100
66
72
67
87
85
95
71
68
51
74
83
86
88
92
100
54
84
85
99
71
68
95
90
100
98
97
99
100
100
100
100
100
65
92
95
1.00
100
100
100
100
100
100
99
73
100
69
100
100
76
98
100
79
82
99
76
North America
South America
Africa & Arabia
Australia & Pacific
Southeast Asia
Europe
Fig. 1B
ZOYSIEAE
CYNODONTEAE
ZOYSIINAE
A
B
C
D
OUTGROUP
TRIRAPHIDEAE
ERAGROSTIDEAE
CENTROPODIEAE
Sporobolus
SPOROBOLINAE
X
sect. Sporobolus
sect. Fimbriatae
subsect. Crypsis
subsect. Helvoli
subsect. Helvoli
sect. Crypsis
sect. Triachyrum
Eragrostis megalosperma Blake 6966
Eragrostis megalosperma Lazarides 4215
Eragrostis megalosperma Lazarides 5647
1.00
97
1.00
87


Sporobolus: phylogeny and classification
TAX ON

0.1
Sporobolus tourneuxii
Sporobolus pungens Durandi 1846
Sporobolus pungens Zohary 489
Sporobolus virginicus Peterson 14311
Sporobolus virginicus Peterson 15683
Sporobolus virginicus Whistler 6132
Sporobolus virginicus Peterson 23820
Sporobolus virginicus Baldini s.n.
Sporobolus kentrophyllus Bogdan 3306
Sporobolus kentrophyllus Mwasumbi 13049
Sporobolus rigidifolius
Sporobolus verdcourtii
Sporobolus uniglumis
Sporobolus scabriflorus
Sporobolus microprotus
Sporobolus spicatus Peterson 24230
Sporobolus spicatus Laegaard 17790
Sporobolus spicatus Peterson 24055
Sporobolus phleoides
Sporobolus nitens
Sporobolus australasicus Peterson 14404
Sporobolus australasicus Walsh 4237
Sporobolus caroli Saarela 1626
Sporobolus caroli Speak 1915
Sporobolus actinocladus Senaratne 6082
Sporobolus actinocladus Saarela 1670
Sporobolus actinocladus Batianoff s.n.
Sporobolus actinocladus Saarela 1625
Sporobolus centrifugus
Sporobolus cordofanus Laegaard 15973
Sporobolus marginatus Rathay 664
Sporobolus scabridus
Sporobolus arabicus
Sporobolus ludwigii
Sporobolus tenacissimus
Sporobolus ioclados
Sporobolus pyramidatus Peterson 24868
Sporobolus coahuilensis Gonzales 3600
Sporobolus pyramidatus Peterson 8920
Sporobolus contractus
Sporobolus pyramidatus Gooding 191
Sporobolus coromandelianus Fosberg 51193
Sporobolus domingensis
Sporobolus pyramidatus Peterson 21163
Sporobolus pyramidatus Garcia 3757
Sporobolus pyramidatus Peterson 18994
Sporobolus coahuilensis Peterson 10000
Sporobolus marginatus Leippert 5101
Sporobolus coromandelianus Schweinfurth 896
Sporobolus cordofanus Peterson 24232
Sporobolus cordofanus Greenway 10191
Sporobolus coromandelianus Peterson 24269
Sporobolus buckleyi Lira 546
Sporobolus buckleyi Piedra s.n.
Sporobolus buckleyi Rodriguez s.n.
Sporobolus palmeri Peterson 24862
Sporobolus palmeri Peterson 24955b
Sporobolus palmeri Peterson 24918
Sporobolus palmeri Peterson 24955a
Sporobolus splendens
Sporobolus spiciformis Garcia 2638
Sporobolus spiciformis Garcia 2814
Sporobolus wrightii Peterson 10638
Sporobolus airoides subsp. airoides Peterson 24853
Sporobolus airoides subsp. airoides Peterson 24587
Sporobolus airoides Peterson 24895
Sporobolus wrightii Peterson 24841
Sporobolus airoides Peterson 10002
Sporobolus wrightii Peterson 19841
Sporobolus airoides Peterson 24956
Sporobolus texanus
Sporobolus nealleyi Peterson 17839
Sporobolus nealleyi Villarreal 1991
Sporobolus giganteus
Sporobolus flexuosus Valdes Reyna 2014
Sporobolus cryptandrus Peterson 24454
Sporobolus flexuosus Reeder 5477
Sporobolus cryptandrus Peterson 24485
Sporobolus cryptandrus Peterson 22003
Sporobolus rigens
Sporobolus compositus
Sporobolus clandestinus Waterfall 5881
Sporobolus neglectus
Sporobolus clandestinus Freeman 6687
Sporobolus clandestinus Waterfall 12309
Sporobolus clandestinus Schuster 197
Sporobolus vaginiflorus Rogers 40059
Sporobolus vaginiflorus Wherry s.n.
Calamovilfa brevipilis
Calamovilfa arcuata
Calamovilfa longifolia Hatch 5738
Calamovilfa gigantea Long 1124
Calamovilfa gigantea Gates 17021
Calamovilfa gigantea Ungor 1018
Sporobolus silveanus
Sporobolus heterolepis
Sporobolus pinetorum
Sporobolus teretifolius McDonald 9988
Sporobolus teretifolius Peterson 14232
Spartina spartinae Reeder 4568
Spartina spartinae Villarreal 3201
Spartina foliosa
Spartina alterniflora Lakela 26573
Spartina alterniflora Naylov 236
Spartina anglica Matthews s.n.
Spartina anglica Williams s.n. a
Spartina anglica Williams s.n. b
Spartina maritima
Spartina ×townsendii
Spartina gracilis Lewis 78
Spartina gracilis Scoggan 15626
Spartina cynosuroides Fisher 33123
Spartina cynosuroides Hill 15630
Spartina gracilis Hendrickson 41
Spartina ×caespitosa
Spartina pectinata Cooperrider s.n.
Spartina pectinata Dirig 2812
Spartina bakeri
Spartina patens Shchepanek 6426
Spartina patens Dutton 2536
Spartina patens Peterson 24435
Spartina ciliata
Spartina densiflora Peterson 19154
Spartina densiflora Lomer 5723
Spartina montevidensis Clayton 4723
1.00
1.00
1.00
1.00
1.00
0.88
1.00
1.00
1.00
0.95
0.99
0.98
1.00
1.00
0.89
1.00
1.00
0.61
0.51
0.98
1.00
1.00
0.52
1.00
0.98
1.00
1.00
0.81
0.99
0.96
1.00
1.00
0.91
0.98
1.00
0.69
1.00
1.00
1.00
0.86
1.00
0.69
1.00
1.00
1.00
1.00
1.00
0.98
0.80
0.99
1.00
0.74
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.90
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.99
0.89
0.97 1.00
1.00
1.00
0.91
1.00
1.00
87
70
66
100
87
96
99
64
100
100
100
100
100
84
64 92
78
99
72
54
100
100
60
100
89
96
62
72
67
100
81
64
63
61
85
91
67
100
97
87
100
83 75
68
0.80
53
52
100
100
76
98
99
77
53
98
97
100
100
99
99
98
98
72
100
67
100
96
100
93 56
91
53
100
100
72
100
100
75
100
100
North America
South America
Africa & Arabia
Australia & Pacific
Southeastern Asia
Europe
Fig. 1A
E
F subsect. Subulati
G
H
I
J
K
L
M
N
O
North American clade
subsect. Ponceletia
subsect. Alterniflori
subsect. Spartina
sect. Spartina
subsect. Floridani
subsect. Calamovilfa
sect. Calamovilfa
sect. Clandestini
sect. Cryptandri
sect. Airoides
sect. Virginicae
sect. Pyramidati
subsect. Spicati
subsect. Actinocladi
subsect. Pyramidati
Sporobolus vaginiflorus var. ozarkanus
Sporobolus vaginiflorus Wieboldt 9596
0.60
0.99
1.00
77
97
Fig. 1A–B.







 




TAX ON


Sporobolus: phylogeny and classification
Version of Record (identical to print version).
Gymnopogon grandiflorus
Leptothrium rigidum
Eleusine indica
Chloris barbata
Cynodon indicus
Pogononeura biflora
Melanocenchris abyssinica
Eragrostiella leioptera Oropetium capense
Tripogon trifidus
Psilolemma jaegeri Peterson 24247
Psilolemma jaegeri Peterson 24249
Sporobolus somalensis Boalev 17
Sporobolus somalensis Herruming 2022
Urochondra setulosa Inckennon 181
Urochondra setulosa Rechinger 27496
Urochondra setulosa Bailey10868
Urochondra setulosa Baldini s.n.
Zoysia pacifica Lorence 9432
Zoysia pacifica Lorence 7651
Zoysia macrantha
Zoysia japonica
Zoysia macrantha walshii
Sporobolus tenellus
Sporobolus acinifolius Smook 3530
Sporobolus acinifolius Smook 10167
Sporobolus albicans Smook 2459
Sporobolus albicans Smook 6270
Sporobolus scabriflorus
Sporobolus farinosus
Sporobolus tourneuxii
Sporobolus mildbraedii
Sporobolus dinklagei
Sporobolus purpurascens Trovart 25
Sporobolus purpurascens Peterson 9453
Sporobolus purpurascens Swallen 10179
Sporobolus acuminatus Guala 1372
Sporobolus linearifolius
Sporobolus acuminatus Irwin 11586
Sporobolus aeneus
Sporobolus apiculatus
Sporobolus eylesii
Sporobolus junceus Strong 2332
Sporobolus junceus Thierat 25181
Sporobolus macrospermus Peterson 9857
Sporobolus lasiophyllus Peterson 21879
Sporobolus lasiophyllus Peterson 21870
Sporobolus bogotensis
Sporobolus lasiophyllus Peterson 21820
Sporobolus sanguineus Bidgood 2397
Sporobolus sanguineus Gereau 6014
Sporobolus panicoides Bidgood 1005
Sporobolus panicoides Smook 9865
Sporobolus micranthus
Sporobolus subglobosus
Sporobolus pilifer
Sporobolus stolzii Peterson 24133
Sporobolus stolzii Peterson 23946
Sporobolus stolzii Richards 21377
Sporobolus fimbriatus Peterson 24206
Sporobolus greenwayi
Sporobolus fimbriatus Peterson 24241
Sporobolus fimbriatus Peterson 24280
Sporobolus diffusus
Sporobolus confinis
Sporobolus brockmanii
Sporobolus nervosus
Sporobolus phyllotrichus
Sporobolus pellucidus
Sporobolus maderaspatanus
Sporobolus agrostoides
Sporobolus montanus
Sporobolus virginicus Peterson 15683
Sporobolus pungens Zohary 489
Sporobolus virginicus Peterson 14311
Sporobolus oxylepsis
Sporobolus robustus
Sporobolus virginicus Baldini s.n.
Sporobolus humilis
Sporobolus virginicus Peterson 23820
Sporobolus virginicus Whistler 6132
Sporobolus tenuissimus Greenway 1745
Sporobolus atrovirens Peterson 24661
Sporobolus molleri Gereau 5790
Sporobolus molleri Rwaburindore 2183
Sporobolus pectinellus Peterson 23978
Sporobolus diandrus
Sporobolus blakei
Sporobolus tenacissimus
Sporobolus tenuissimus Peterson 9523
Sporobolus trichodes
Sporobolus atrovirens Peterson 22342
Sporobolus atrovirens Peterson 24729
Sporobolus atrovirens Rosalen 3381
Sporobolus pyramidalis Peterson 24150
Sporobolus pyramidalis Senaratne 6082
Sporobolus stapfianus
Sporobolus pectinellus Fay 7131
Sporobolus festivus
Sporobolus infirmus
Sporobolus myrianthus
Sporobolus sessilis
Sporobolus africanus Peterson 24024
Sporobolus africanus Peterson 24121
Sporobolus fertilis Raulerson 775
Sporobolus fertilis Gould 13535
Sporobolus natalensis
Sporobolus creber
Sporobolus laxus
Sporobolus pseudairoides
Sporobolus indicus Peterson 22025
Sporobolus berteroanus
Sporobolus indicus Peterson 7337
Sporobolus jacquemonthii Estrada 18964
Sporobolus jacquemontii Peterson 15902
North America
South America
Africa & Arabia
Australia & Pacific
Southeast Asia
Europe
1.00
100
1.00
100
1.00
100 1.00
100
1.00
100 1.00
100
1.00
100
1.00
100
1.00
100
1.00
100
1.00
100
1.00
88
1.00
100
1.00
0.96
1.00
100
1.00
100
1.00
100
1.00
0.71
1.00
0.95
0.93
1.00
0.99
1.00
1.00
1.00
1.00
1.00
1.00
1.00 1.00
0.99
1.00
1.00
1.00
1.00
1.00
0.96
1.00
1.00
0.92
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.80
0.59
1.00
1.00
1.00
1.00
1.00
1.00
0.99
1.00
1.00
0.82
0.53
1.00
0.99
0.64
1.00
1.00
1.00
OUTGROUP
TRIRAPHIDEAE
ERAGROSTIDEAE
CENTROPODIEAE
1.00
ZOYSIEAE
CYNODONTEAE
ZOYSIINAE
sect. Sporobolus
B
sect. Triachyrum
E
95
51
51
100
100
51
63
96
100
100
100
100
100
96
100
96
71
53
100
59
54
60
99
96
88
1.00
79
86
76
100
96
97
97
100
96
62
81
100
100
1.00
100
1.00
87
97
97
100
74
65
100
62
100
99
93
100
Fig. 2B
0.2
Sporobolus
91
52
1.00
96
59
54
89
99
100
X
A
D
sect. Virginicae
sect.
Fimbriatae
Fig. 2A–B.
  




Sporobolus: phylogeny and classification
TAX ON

0.2
Sporobolus microprotus
Sporobolus uniglumis
Sporobolus spicatus Peterson 24055
Sporobolus spicatus Laegaard 17790
Sporobolus spicatus Peterson 24230
Sporobolus nitens
Sporobolus olivaceus
Sporobolus phleoides
Sporobolus australasicus Peterson 14404
Sporobolus australasicus Walsh 4237
Sporobolus actinocladus Senaratne 6082
Sporobolus caroli Speak 1915
Sporobolus actinocladus Saarela 1625
Sporobolus caroli Saarela 1626
Sporobolus scabridus
Sporobolus actinocladus Batianoff s.n.
Sporobolus actinocladus Saarela1670
Sporobolus ludwigii
Sporobolus subulatus
Sporobolus verdcourtii
Sporobolus arabicus
Sporobolus kentrophyllus Bogdan 3306
Sporobolus kentrophyllus Mwasumbi 13049
Sporobolus domingensis
Sporobolus ioclados
Sporobolus centrifugus
Sporobolus cordofanus Laegaard 15973
Sporobolus marginatus Rathay 664
Sporobolus marginatus Leippert 5101
Sporobolus coromandelianus Peterson 24269
Sporobolus coromandelianus Schweinfurth 896
Sporobolus cordofanus Greenway 10191
Sporobolus cordofanus Peterson 24232
Sporobolus pyramidatus Peterson 24868
Sporobolus contractus
Sporobolus coahuilensis Gonzales 3600
Sporobolus coahuilensis Peterson 10000
Sporobolus pyramidatus Gooding 191
Sporobolus pyramidatus Peterson 18994
Sporobolus pyramidatus Garcia 3757
Sporobolus coromandelianus Fosberg 51193
Sporobolus pyramidatus Peterson 8920
Sporobolus pyramidatus Peterson 21163
Sporobolus neglectus
Sporobolus consimilis Collenette 17
Sporobolus consimilis Peterson 24252
Crypsis aculeata Soreng 7940
Crypsis alopecuroides
Crypsis aculeata Soreng 5469
Crypsis schoenoides
Sporobolus ruspolianus Bally 15581
Sporobolus ruspolianus McKinnon s.n.
Sporobolus ruspolianus Flemming 2136
Sporobolus mitchellii
Sporobolus helvolus Laegaard 17063
Sporobolus helvolus Peterson 24217
Sporobolus elongatus
Sporobolus palmeri Peterson 24918
Sporobolus palmeri Peterson 24955b
Sporobolus palmeri Peterson 24862
Sporobolus palmeri Peterson 24955a
Sporobolus buckleyi Lira 546
Sporobolus buckleyi Piedra s.n.
Sporobolus buckleyi Rodriguez s.n.
Thellungia advena Belson 1930
Thellungia advena Lazarides 4185
Sporobolus compositus
Sporobolus conrathii
Sporobolus aldabrensis
Sporobolus clandestinus Freeman 6687
Sporobolus clandestinus Schuster 197
Sporobolus clandestinus Waterfall 12309
Sporobolus clandestinus Waterfall 5881
Sporobolus clandestinus Peterson 24422
Sporobolus vaginiflorus Rogers 40059
Sporobolus vaginiflorus Wherry s.n.
Sporobolus texanus
Sporobolus nealleyi Correll 18548
Sporobolus nealleyi Peterson 17839
Sporobolus nealleyi Villarreal 1991
Sporobolus cryptandrus Peterson 24454
Sporobolus cryptandrus Peterson 24485
Sporobolus flexuosus Reeder 5477
Sporobolus flexuosus Valdes Reyna 2014
Sporobolus giganteus
Sporobolus airoides Peterson 24895
Sporobolus airoides subsp. airoides Peterson 24587
Sporobolus airoides Peterson 24956
Sporobolus wrightii Peterson 10638
Sporobolus wrightii Peterson 19841
Sporobolus rigens
Sporobolus airoides subsp. airoides Peterson 24853
Sporobolus spiciformis Garcia 2814
Sporobolus spiciformis Garcia 2638
Sporobolus airoides Peterson 10002
Sporobolus wrightii Peterson 24841
Calamovilfa arcuata
Calamovilfa brevipilis
Calamovilfa gigantea Long 1124
Calamovilfa curtissii
Calamovilfa longifolia
Calamovilfa gigantea Gates 17021
Calamovilfa gigantea Ungor 1018
Sporobolus teretifolius Peterson 14232
Sporobolus floridanus Harper s.n.
Sporobolus pinetorum
Sporobolus teretifolius McDonald 9988
Sporobolus floridanus Curtiss s.n.
Sporobolus heterolepis Davidse 19101
Sporobolus silveanus
Spartina maritima Fernandez Casas 5537
Spartina maritima Marchant s.n.
Spartina foliosa
Spartina alterniflora Naylov 236
Spartina alterniflora Lakela 26573
Spartina anglica Williams s.n. a
Spartina ×townsendii Saarela 791
Spartina anglica Matthews s.n.
Spartina anglica Williams s,n. b
Spartina ×townsendii Hubbard s.n.
Spartina spartinae Reeder 4568
Spartina spartinae Villarreal 3201
Spartina densiflora Peterson 19154
Spartina densiflora Lomer 5723
Spartina montevidensis Clayton 4723
Spartina cynosuroides Fisher 33123
Spartina cynosuroides Hill 5630
Spartina ciliata Rambo 56450
Spartina montevidensis Rosengurtt 10853
Spartina gracilis Lewis 1013
Spartina gracilis Scoggan 15626
Spartina patens Shchepanek 6426
Spartina patens Dutton 2536
Spartina bakeri
Spartina patens Peterson 24435
Spartina gracilis Hendrickson s.n.
Spartina ×caespitosa Eaton 587
Spartina pectinata Cooperrider s.n.
North America
South America
Africa & Arabia
Australia & Pacific
Southeast Asia
Europe
1.00
1.00
1.00
1.00
1.00
1.00
0.97
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.51
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.84
1.00
1.00
0.98
0.97
0.72
0.56 0.91
1.00
0.96
0.63
0.67
0.67
0.91
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
1.00
0.69
C
F
G
H
I
J
K
L
M
N
O
63
99
96
100
100
100
100
94
87
100
96
100
100
99
76
97
100
100
90
90
54
100
62
69
99
94
93
74
69
61
100
100
100
100
54
93
100
1.00
79
1.00
100
1.00
1.00
1.00
100
100
100
100
100
100
90
100
1.00
75
93
Fig. 2A
North American clade
98
sect. Spartina
subsect. Alterniflori
subsect. Ponceletia
subsect. Spartina
subsect. Floridani
subsect. Calamovilfa
sect. Calamovilfa
sect. Clandestini
sect. Airoides
sect. Cryptandri
subsect. Crypsis
subsect. helvoli sect.
sect. Pyramidati
subsect. Spicati
subsect. Actinocladi
subsect. Subulati
subsect. Pyramidati
Sporobolus vaginiflorus Peterson 24441
Sporobolus vaginiflorus var. ozarkanus
Sporobolus splendens
1.00
91
Eragrostis megalosperma Blake 6966
Eragrostis megalosperma Lazarides 4215
Eragrostis megalosperma Lazarides 5647
1.00
99
1.00
94

TAX ON


Sporobolus: phylogeny and classification
Version of Record (identical to print version).



  


Spartina alternif lora S. anglica S. foli
osa S. maritima 
   S. ×caespitosa S. ciliata S. cynosuroi
des S. densi flora S. gracilis   S. patens
S. pectinata 





Sporobolus neglectus Sporobolus
consimilisSporobolus elongatus
S. palmeri Eragrostis megalo
sperma 

Incongruences between the ITS and combined plastid
phylograms. —
 
 





Sporobolus
arabicusS. humilisS. scabridus
S. scabriflorus
niflorus 
 S. conrathii 
S. neglectusS. rigensS. robustus
EragrostismegalospermaS. S. con
Table 2.
Sporobolus
boldAOX


 Sporobolus somalensis
Psilolemma P. jaegeriP. jaegeri
SporobolusS. indicus
S. acinifolius  ThellungiaS. albicans nn
S. conrathii  S. consimilis  S. elongatus n S. megalospermus 
 S. palmeri  S. rigens  S. tenellus 
S.S. airoidesJS. airoides n S. spiciformis n
S. splendens .n
S. M, N
MS. arcuatus  S. arenicola 
tean S. curtissianus  S. rigidus n

S. floridanusN*S. curtissii  S. floridanus S. heterolepis
n *S. interruptus n S. pinetorum  S. teretifolius

S.ClandestiniS. clandestinusLS. aldabrensis S. compositus n
 S. clandestinus S. neglectus nn
S.CrypsisS. aculeatusC
CrypsisCS. aculeatus S. alopecuroides n
 *S. minuarti
oides  *S. niliacus n S. schoenoides 
n
Cn S. mitchellii 
 S. ruspolianus 
S.  Cryptandri S. cryptandrusKS. cryptandrus n
nS. giganteusnS. nealleyin
S.S. fimbriatus BS. agrostoides  S. confinis 
S. diffusus  S. fimbriatus n*S. macranthus  S. maderaspatanus
n S. montanus nn S. pellucidus  S. phyllotrichus 
S. smutzii 
S.  Pyramidati S. pyramidatusF, G, H, I
Pyramidati IS. centrifugus  S. coahuilensis  S. contractus n
S. cordofanus  S. coromandelianus n S. domingensis  S. ioclados
n S S. marginatus n S. pyramidatus n
 S. scabridus S. tenacissimus 


Sporobolus: phylogeny and classification
TAX ON

Version of Record (identical to print version).
similisS. elongatusS. palmeriS. somalensisS. 



DISCUSSION

Sporobolus  Cryp
sis Spartina  ThellungiaPsi
lolemma jaegeriSporobolus somalensis   
SporobolusSporobolus




Sporobolus




  

 Crypsis Spartina
ThellungiaSporobolus


 
     
Sporobolus. 
 Sporobolus




S. actinocladusHS. actinocladus S. australasicus  S. caroli 
*S. contiguus  S. nitens *S. partimpatens 
S. phleoides  *S. pulchellus 
SpicatiS. spicatusGS. microprotus  S. scabriflorus  S. spicatus
S. uniglumis
SubulatiS. subulatusFS. arabicus n
n S. subulatus
S.SpartinaS. cynosuroidesO
alterniflorusS. alterniflorus n
S. anglicus nS. foliosusn
S. ×longispicusS. martimusn
n
PonceletiaS. mobberleyanusSpartina arundinacea
S. mobberleyanusn S. spartinus n
Spartina S.n S. coarctatus
Spartina ciliataS. cynosuroides n S. densiflorus
S. ×eatonianusSpartina ×caespitosa n
nusSpartina gracilisnSpartina pectinta
nS. pumilusSpartina patensn
S.SporobolusTruncataeS.
S. indicusAS. africanus nn S. berteroanus
 S. creber  S. diandrus n S. farinosus 
S. fertilis nn *S. fourcadii  S. indicus n
S. infirmus  S. jacquemontii n*S. minor  S. mollerin S. natalensis
 S. myrianthus  S. pectinellus  S. pseudairoides  S. pyramidalis n
*S. quadratus  S. sessilis  S. stapfianus  S. tenuissimus n S. trichodes 
S.TriachyrumS. discosporusDS. acuminatus  *S. adustus 
 S. aeneus  *S. amaliae  S. apiculatus  S. bogotensis 
 *S. camporum *S. cubensis *S. discosporus
S. eylesii  *S. harmandii  S. junceus  S. lasiophyllus  S. linearifolius
 S. macropsermus  S. micranthus  S. mildbraedii  *S. multinodis 
 *S. nudiramus . panicoides  *S. paniculatus 
*S. paucifolius  S. pilifer n S. purpurascens n
 S. sanguineus n *S. sciadocladus
 S. stolzii  S. subglobosus 
S.VirginicaeES. humilis n S. pungens  S. robustus 
n
Table 2.


TAX ON


Sporobolus: phylogeny and classification
Version of Record (identical to print version).
Cynodonteae. —
Pogononeura biflora 

 
Sporobolus 

    
P. biflora 
Sporobolus cordo
fanus  
 
P. bifloraS. cordofanus

Turner & Watson

 P. bif lora 
P. biflora

Sporobolinae and Sporobolus, incertae sedis. —



A
B
C
D
E
F
G
H
I
J
K
L
M
N
O
H
I
M
O
B
C
D
E
F
G
J
K
L
N
Thellungia advena
Sporobolus oxylepsis
Sporobolus robustus
Sporobolus humilis
Sporobolus arabicus
North American clade
North American clade
Afr
Afr
Afr Afr
Afr Afr
Aus
Aus
XX
S.Afr
S.Afr
Sporobolus somalensis
Psilolemma
Urochondra
Zoysia
Sporobolus somalensis
Psilolemma
Urochondra
Zoysia
Calamovilfa clade
Spartina clade
Sporobolus sect. Crypsis clade
ITS Plastid
(rpl32-trnL, ndhA, rps16, rps16-trnK)
Sporobolus floridanus clade
Sporobolus compositus clade
Sporobolus pyramidatus clade
(F, G. H, I)
Sporobolus virginicus clade
Sporobolus junceus
clade
Sporobolus fimbriatus
clade
Sporobolus indicus clade
Sporobolus acinifolius–S. albicans–S. tenellus
Sporobolus scabridus
Sporobolus airoides clade
Sporobolus cryptandrus clade
Eragrostis megalosperma
Sporobolus consimilis
Sporobolus rigens
A
Afr
Aus
Afr
Aus
Afr
Aus
Afr
Aus
Sporobolus vaginiflorus var. ozarkanus
Sporobolus neglectus
Sporobolus scabriflorus
Fig. 3.
Sporobolus