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This is the first phylogenetic study focused on the Pavetteae, one of the most species-rich and morphologically diverse tribes within the coffee family (Rubiaceae). Fifteen of the 17 currently recognized genera, represented by 85 taxa, were sequenced for rps16, trnT-F and ITS and analysed using Bayesian inference and maximum likelihood methods. The monophyly of the Pavetteae is confirmed. Four major lineages are identified, but their phylogenetic relationships are not fully resolved. The continental African genera Rutidea, Nichallea and Tennantia, the Madagascan genera Homollea and Robbrechtia, and the paleotropical genus Pavetta are monophyletic. Other genera are paraphyletic in their current circumscriptions and the following changes are made: Homolliella is placed in synonymy with Paracephaelis, and Coleactina and Dictyandra with Leptactina, resulting in four new combinations. The large paleotropical genus Tarenna is shown not to be monophyletic. In the future, the name Tarenna should not be used for continental African species. Most of these could be transferred to the hitherto monospecific genus Cladoceras, but other species might constitute altogether new genera. The relationship between the monophyletic Asian-Pacific and Madagascan Tarenna species remains unclear. The phylogeny of the Madagascan genera of the Pavetteae is largely unresolved and the largest Madagascar-centred genus Coptosperma was not recovered as monophyletic. The low resolution for the Madagascan taxa can be considered as an indication of rapid radiation. Further molecular and morphological studies are necessary to clarify the phylogeny of the Pavetteae, especially regarding the African Tarenna species and the Madagascan genera of the tribe.
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De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
TAX O N
 
Version of Record

;   || 
INTRODUCTION
  

-
 Pavetta   Ta r e n n a
Pachy-
stylusTr if l o-
rensia
          
-

   
    


   
    
  Ixora Pavetta  


    
    



 Tar e n n aLeptactina -
 
-
 

     


Molecular phylogenetics and generic assessment in the tribe Pavetteae
(Rubiaceae)
Petra De Block,1 Sylvain G. Razafimandimbison,2 Steven Janssens,1 Helga Ochoterena,3 Elmar Robbrecht1
& Birgitta Bremer2
 
  
 
 

DOI
 
Abstract
            


    
Nichallea Te n n antia  
Pavetta
 ParacephaelisLeptactina
Ta renn a  
Tare n n a-
 
Tare n n a  
  
-
Taren n a 

Keywords
;  ; Tare n n a
Supplementary Material

 
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TAX O N
 De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
Version of Record

    -


Tarenna
    -
 
 Ixora
          


Ixora
 
   
-
    -
     
 
Ixora
     
Tr i-
florensia  
          -

Table 1.

 

 




  1  
Tarenna
 N.Hallé  1 

= 


19   


 2 
  3  
  1 
Leptactina  ca. 20   Leptactina 

Nichallea  1  Nichallea
Pachystylus  2  Pachystylus
Paracephaelis Baill.    Paracephaelis 

Pavetta L.     Pavetta
 De Block  2  
  22   


    
TarennaB  ca. 200   Tarenna

Tennantia  1  Tennantia
Triflorensia

 3

7Triflorensia

Ixora


-

Tarenna
81
De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
TAX O N
 
Version of Record
Fig. 1.

A,

B,

C,
Paracephaelis cinerea

D,
Paracephaelis tiliacea 
E,
 
F,
 
G,


H,

I,

J,

  
K,
  
L,
  
M,
Tarenna thouarsiana  
N,
 
rescens
O,
Leptactina 
P,


82
TAX O N
 De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
Version of Record



   


Tare n n a    -


     

   -
            


-


MATERIALS AND METHODS
Taxon sampling. —
       
 Pachystylus  Triflorensia
      
 

Nichallea   Tennantia   
    
  -
  
Pavetta   
  





-
actinaNichallea -



   

Pavetta 

 Pavetta -
     -
  


DNA isolation, amplification and sequencing. —
   
  
         
-


-
     

   




  


-

  

TaqTaq

  
-
 
    
   
     

Table 2.

   
rps16   
 
trnT-trnL   
 
trnL-   
 

 


 
 
 
83
De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
TAX O N
 
Version of Record
Sequence alignment and phylogenetic analyses. —
   
-




   
    

& al.


  -


  


          
   

   

  

    

-

RESULTS
  

      .

   Para-
cephaelis saxatilis






    
    
    
 
 -
   

  
    
NichalleaLeptactina  



-


-

         -



      -
Nichallea   

 
-



-
Leptactina
Leptactina

papalis Leptactina 
  
      


= Leptactina.
      -
Pavetta
Tar e n n a 
Tarenna jolinonii
    
 

 -


Ta r e n n a 
  
Pavetta-
sis
Pavetta
Pavetta
Table 3.

rps16   
  93 79 
 918 2217  
 738 1777  3129
 180  281 901
 117 293  
 12.7 13.2 12.8 13.0
  30 89

TAX O N
 De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
Version of Record
Razafimandimbisonia humblotii
Mussaenda flava
Vangueria madagascariensis
Ixora sp.
Tricalysia semidecidua
Oxyanthus troupinii
Gardenia rutenbergiana
Euclinia longiflora
Nichallea soyauxii
Rutidea membranacea
Rutidea fuscescens
Rutidea orientalis
Rutidea parviflora
Rutidea seretii
Rutidea olenotricha
Rutidea dupuisii
Rutidea ferruginea
Rutidea decorticata
Leptactina leopoldisecundi
Leptactina involucrata
Dictyandra arborescens
Leptactina mannii
Leptactina pynaertii
Leptactina delagoensis
Leptactina papyrophloea
Leptactina epinyctios
Leptactina benguelensis
Coleactina papalis
Tarenna jolinonii
Tarenna fuscoflava
Tarenna lasiorachis
Tarenna nitidula
Tarenna pallidula
Tarenna vignei
Tarenna roseicosta
Cladoceras subcapitatum
Tarenna pembensis
Tarenna precidantenna
Tarenna bipindensis
Pavetta tetramera
Pavetta suffruticosa
Pavetta ternifolia
Pavetta hymenophylla
Pavetta sp. A (FTEA)
Pavetta tarennoides
Pavetta sansibarica
Pavetta schumanniana
Pavetta stenosepala
Pavetta batesiana
Pavetta vaga
Pavetta agrostiphylla
Pavetta indica
Pavetta sp. B
Pavetta sp. C
Tennantia sennii
Tarenna alpestris
Tarenna asiatica
Tarenna flava
Tarenna leioloba
Tarenna rhypalostigma
Tarenna microcarpa
Tarenna sambucina
Tarenna gracilipes
Tarenna attenuata
Tarenna depauperata
Coptosperma graveolens
Coptosperma peteri
Homollea perrieri
Homollea longiflora
Paracephaelis tiliacea
Homolliella sericea
Paracephaelis cinerea
Paracephaelis saxatilis
Robbrechtia grandifolia
Robbrechtia milleri
Tarenna spiranthera
Tarenna grevei
Tarenna capuroniana
Tarenna alleizettei
Tarenna uniflora
Tarenna thouarsiana
Coptosperma supra-axillare
Coptosperma nigrescens
Coptosperma nigrescens
Coptosperma sp. D
Coptosperma sp. E
Coptosperma madagascariense
Coptosperma sp. A
Coptosperma littorale
Coptosperma sp. B
Coptosperma sp. C
Schizenterospermum rotundifolium
Schizenterospermum grevei
Coptosperma borbonica
Continental
Africa
Australia
Asia/Pacific
Asia
Continental
Africa
Continental
Africa
Madagascar
Madagascar
Continental Africa
Madagascar
Mascarenes
Continental Africa
+ Madagascar
100/1.0
57/0.95
89/1.0
93/1.0
95/1.0
70/1.0
81/1.0 98/1.0
82/1.0
92/1.0
65/0.66 76/1.0
85/0.98
100/1.0
67/0.78
74/1.0
65/0.53
80/0.96
71/0.99
100/1.0
54/0.95 * /0.98 100/1.0
88/1.0
94/1.0
* /0.96
* /0.90
* /0.59
* /0.78
60/0.73
100/1.0
* /0.84
96/1.0
58/0.66
90/0.93
* /0.55
* /0.86
100/1.0
98/1.0
92/1.0
93/1.0
51/0.81
97/1.0
* /0.97
* /0.85
69/0.99
95/1.0
100/1.0
97/1.0
90/0.71
52/0.56
82/0.99
52/0.64
76/1.091/1.0
100/1.0
71/0.79
100/1.0 66/0.93
62/0.99
74/1.0
*/0.51
70/1.0
76/0.95
94/1.0
91/1.0
100/0.94
100/1.0
57/0.54
72/1.0
98/1.0
* /0.65
92/1.0
100/1.0
100/1.0
100/1.0
51/0.50
55/0.91
75/0.97
97/1.0
* /
0.77
100/1.0
100/1.0
*
*
*
*
*
*
*
*
*
**
*
*
*
*
*
Clade II
Clade III
Clade IV
Clade I
Pavetteae
Fig. 2.
-
    
-


 
 

.

De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
TAX O N
 
Version of Record
 

Pavetta  Baconia 
  
Pavetta   

 Pavetta
       
 
 
         


Tennantia Tarenna

Para-
cephaelis
           
   -
   -
Tennantia

Tarenna

   -

Paracephaelis

sericeaParacephaelis
Tare n n a 
    
 
   


    

    
   
  -
 


        
  
   
-
cens  

      
Tare n n a  
-
  Tarenna 
Paracephaelis

Tare n n a Ta r e n n a
Pavetta
DISCUSSION
Monophyly and main lineages of the Pavetteae. —

         

-

    -


      

  

  
    
Nichallea -
andraLeptactina
Ta r e n n a      Pavetta.
   -
-
TennantiaParacephaelis
Tarenna
ParacephaelisTarenna
   TarennaPara-
cephaelis
 
     

     


     
    
      -

-

.
        
      
         


Tennantia
Tarenna 
  
-

 
  

     
    



 -


TAX O N
 De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
Version of Record
Relationships and circumscriptions of the genera of the
Pavetteae. —
      
        -
Nichallea
Pavetta   -
Tennantia
    
    Leptactina
Paracephaelis 
Leptactina
 Tarenna  


Rutidea and Nichallea (clade I). —
  

Nichallea soyauxii
Nichallea. 
Nichallea

      

Nichallea    
   

    
-


       
    -

    
Nichallea    
 Nichallea-
  Nichallea -
    Nichallea  -
 Nichallea 

Nichallea




        
Nichallea

      
 
  -
 

  
    




Coleactina, Dic tyandra and Leptactina (clade II). —

    
Leptactina-
tina papalis.   Leptactina







 -
tina Leptactina


-
actina  Leptactina  

 Leptactina.
   
rescens     Leptactina   
  


Leptactina
  
Leptactina.
     Leptactina
       
      
 Leptactina  
 

  Leptactina

Leptactina
      
   -
 
 
    -
 
     
 
    
     

Leptactina
 
       
 


      
   
    


.
Leptactina         -
       
   
       

Tarenna, Pachystylus, Triflorensia (clades III and IV). —
Tarenna   
   -

87
De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
TAX O N
 
Version of Record
     
Tarenna

  
& al.& al.
  Tarenna  
-
; De Block & al.
  
& al.
Ta r e n n a-

Tarenna
-


   
  
 
Pavetta
 
Tare n n a 
Tare nna.
Tar e n n a   
    -
Ta renna


        
            
 
    

Tare n n a 

   Tarenna


            
-

 Tarenna
         
   

TarennaPachystylusTriflorensia
     
Pachystylus  

TarennaTr if lo -
rensia-
Tar e n n a        


      
 

 
  

Triflorensia.
& al.
 vitiense
TarennaPela-

   

 

           
    & al.



 Tarenna  


-
 thouarsiana

  

 

= 



    
   
Tare n n a     -
sis

 -

  Ta renna

  Pavetta   
ceras       
  
     
Tarenna     

  

Tarenna junodii

     
Tarenna  -
          
   Ta r e n n a 
Leptactina
Pavetta       

88
TAX O N
 De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
Version of Record
Tarenna  

         

   Leptac-
tina


Tarenna. We do
-
Tarenna.
Tarenna      
 Tarenna jolinonii 
Tarenna

III     
Tarenna
    
   -
Tarenna   



;
 



 Tarenna  

Pavetta

Pavetta
    
PavettaPavetta
   
       -
Tarenna   
 Pavetta    
Tar e n n a Pavetta 
  


-
Tarenna

Tarenna Pavetta.
   -

     


      Pavetta
Tarenna -
   
    
 
Pavetta
    
   
Tarenna Pavetta
Tare n n a   Pavetta-

Tarenna-
.
Tarenna


 Tarenna
          

    
   

Pavetta (clade III). —
Pavetta   


Pavetta 
   


     

Pavetta         

PavettaBaconia-
    

     -
      
Pavetta



Pavetta
Baconia  
 Baconia   
Pavetta
     Pavetta
-
phyllaPavetta
 Baconia

Pavetta-

Baconia-
phyllaPavetta    

      Pavetta.
   

    
        
   
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De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
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 
Version of Record
  
Baconia
   

Baconia
Baconia         -
  



  Pavetta  Baconia-


  Pavetta  
 -
     

 Pavetta -
  
 
Pavetta-

-
Pavetta
  
Pavetta       

   Baconia  Pavetta
  
.
Tennantia (clade IV). —

    

  Tarenna  
Tennantia sennii -
Tarenna  




 Tennantia
 Ta renna   

  
TennantiaTare n n a 
    Tennantia   
Tarenna 
Tennantia
Ta r e n n a
Tennantia

    

 Tarenna      

 
TennantiaTar e n n a 
Homollea, Homolliella and Paracephaelis (clade IV). —

    -
   Paracephaelis
      

   -
 


 
PavettaPavetta
-
Ta r e n n a



 Paracephaelis     
-

 

Paracephaelis

Paracephaelis-
   
   
 Paracephaelis         
 -
  Paracephaelis
 Ta r e n n a 
Tarenna
 Paracephaelis

Paracephaelis
Tare n n a 

Paracephaelis
 


 
     
Para-
cephaelis
Paracephaelis.
     Paracephaelis-
-

 
-
lea
Paracephaelis

Coptosperma and Schizenterospermum (clade IV). —
   & al.
  & al.       

90
TAX O N
 De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
Version of Record
Tare n n a        
-
-





 
         
 
-

-
      

-
                        
    
  
      


 

-
-

Tar e n n a -


     
-

  
Tarenna-

      -
 Ta r e n n a 
& al.
-


  

    
   
   
    


-
olens     
 clade.
 

  
         
 
     -
 

         
 
       

 -
 

  
-
  
  
  
 
          
  -




    
   
axillare 


 
-

    
  
     

   -




      
 -


        -
cariense     
Baill     -
    
         


 


   
       
91
De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
TAX O N
 
Version of Record
-
  
-
          -
-

    


    



 -


 

Robbrechtia (clade IV). —
   -

   

-

-

 -


-
.


-
  sericea    

 

NOMENCLATURAL CHANGES

     -

Leptactina   


= 
sy n. n ov.

=
syn. nov.  N.Hallé.
 
  


  
    
-
-
                          
-
-





-






 

-


-

-


Leptactina arborescens
 comb. nov.


Leptactina congolana  comb. nov. -



Leptactina papalis   comb. nov.-
tina papalis
-
Le

ParacephaelisPara-
cephaelis tiliacea Baill.
= syn.
nov.
    
-


 -

 
92
TAX O N
 De Block & al. • Molecular phylogenetics of Pavetteae (Rubiaceae)
Version of Record
Ali, S.J. & Robbrecht, E.    
Tricalysia

Andreasen, K.  -
        
Andreasen, K. & Bremer, B. 

Andreasen, K. & Bremer, B. 
             

 
Arènes, J. 

Baillon, H.N. 

Bremekamp, C.E.B.        Pavetta L.

Bremekamp, C.E.B.      Pavetta 
     

Bremekamp, C.E.B. Pavetta -
   
Bremekamp, C.E.B. .  

Bremer, B. & Eriksson, T.  
   
  
Bridson, D.M.           Kew

Bridson, D.M. Nichallea 
   
Bridson, D.M. Tare n n a-
            
 
    
Bridson, D.M.  Ta ren n a      


Bridson, D.M. Pavetta
     
    
Bridson, D.M.     

Bridson, D.M.  

 
Bridson, D.M. & Robbrecht, E.        

 
Candolle, A.P. de 

Capuron, R.        
   
         
            

Cortes-B., R., Delprete, P.G. & Motley, T.J.  
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