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Page last updated
Thu 17 July 2025

The Freshwater Mussels (Unionoida) of the World (and other less consequential bivalves)

FM(U)otW(aolcb) is the web version of the MUSSEL Project Database. Follow the links to browse the data or use the custom Google search field. Either way, you win!


Bivalvia | Unionoida | Unionoidea | Unionidae | Gonideinae

tribe CONTRADENTINI Modell, 1942: cladomics
     6 genera

Cladomics Summary

number of species = 35 (35 total)
number of subtaxa (rank: genus) = 6
analyzed in 70 cladograms in 44 publications
maximum number of terminals in an analysis = 23 (mean = 6.41, median = 4)
maximum number of subtaxa (genus) = 5 (mean = 2.67, median = 3)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 61
maximum number of included terminals = 23, (mean = 7.21 median = 5)
maximum number of subtaxa (genus) = 5 (mean = 2.92, median = 3)
proportion of cladograms that support basic monophyly = 0.97

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 55
(Rigorous monophyly refers to cladograms with terminal species representing more than one subtaxon (genus) taxon and at least one rigorous outgroup (family).
maximum number of included terminals = 23 (mean = 6.91, median = 4)
maximum number of subtaxa (genus) = 5 (mean = 3, median = 3)
proportion of cladograms that support rigorous monophyly = 0.96


Pairwise Cladogram Clustering

The graphs below depict in two dimensions all pair-wise ingroup distances within the cladome. See the Cladmoics FARQ for more information.


PCoA Ordination


NJ Tree


Pfeiffer et al., 2019 [Publ 3937], fig. 4-5

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
5535521250
  +----------------- Chamberlainiini
  |                                 
  |           +----- Lamprotulini   
--+  +--------+                     
  |  |        |  +-- Gonideini      
  |  |        +--+                  
  +--+           +-- Pseudodontini  
     |                              
     |  +----------- Rectidentini   
     |  |                           
     +--+        +-- Trapezoideus **   
        |  +-----+                  
        |  |     +-- Physunio **       
        +--+                        
           |  +----- Yaukthwa **       
           +--+                     
              |  +-- Lens **           
              +--+                  
                 +-- Solenaia **       

Zieritz et al., 2024 [Publ 4381], fig. 2

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
8864382785
              +-- Schepmaniini 
        +-----+                
        |     +-- Pseudodontini
  +-----+                      
  |     |  +----- Caudiculatus 
  |     +--+                   
  |        |  +-- Lamprotulini 
--+        +--+                
  |           +-- Gonideini    
  |                            
  |  +----------- Ctenodesmini 
  |  |                         
  +--+  +-------- Rectidentini 
     |  |                      
     +--+     +-- Physunio **     
        |  +--+                
        |  |  +-- Trapezoideus ** 
        +--+                   
           |  +-- Yaukthwa **     
           +--+                
              +-- Lens **         

Bolotov et al., 2020 [Publ 4100], fig. S1

iTtTbiriSoTroTroSiD
23232148949522
        +-------- Pseudodontini  
  +-----+                        
  |     |  +----- Gonideini      
  |     +--+                     
  |        |  +-- Chamberlainiini
--+        +--+                  
  |           +-- Lamprotulini   
  |                              
  |  +----------- Rectidentini   
  +--+                           
     |  +-------- Physunio **       
     +--+                        
        |  +----- Trapezoideus **   
        +--+                     
           |  +-- Lens **           
           +--+                  
              +-- Yaukthwa **       

Bolotov et al., 2020 [Publ 4100], fig. 1

iTtTbiriSoTroTroSiD
23232149549520
     +-------- Rectidentini             
     |                                  
  +--+     +-- Trapezoideus **             
  |  |  +--+                            
  |  |  |  +-- Physunio **                 
  |  +--+                               
  |     |  +-- Yaukthwa **                 
  |     +--+                            
  |        +-- Lens **                     
  |                                     
--+     +----- Pseudodontini            
  |     |                               
  |     |  +-- Sinosolenaia_iridinea    
  |     |  |                            
  |     |  +-- Sinosolenaia_recognita__1
  |     |  |                            
  |     |  +-- Sinosolenaia_carinata    
  |     |  |                            
  |     |  +-- Leguminaia               
  +-----+  |                            
        |  +-- Microcondylaea           
        |  |                            
        |  +-- Gonidea                  
        |  |                            
        |  +-- Ptychorhynchus           
        |  |                            
        |  +-- Parvasolenaia            
        |  |                            
        +--+-- Lamprotula_leaii         
           |                            
           +-- Lamprotula               
           |                            
           +-- Lamprotula_caveata       
           |                            
           +-- Potomida                 
           |                            
           +-- Pronodularia             
           |                            
           +-- Sinohyriopsis            
           |                            
           +-- Chamberlainia            
           |                            
           +-- Sinosolenaia_recognita__2
           |                            
           +-- Sinosolenaia_oleivora    

Konopleva et al., 2019 [Publ 4041], fig. 3

iTtTbiriSoTroTroSiD
19191741515218
  +-------------- Pseudodontini
--+                            
  |  +----------- Rectidentini 
  +--+                         
     |  +-------- Yaukthwa **     
     +--+                      
        |  +----- Lens **         
        +--+                   
           |  +-- Trapezoideus ** 
           +--+                
              +-- Physunio **     

Konopleva et al., 2023 [Publ 4262], fig. 8

iTtTbiriSoTroTroSiD
19191746349417
  +-------------- Pseudodontini
  |                            
--+           +-- Gonideini    
  |  +--------+                
  |  |        +-- Lamprotulini 
  +--+                         
     |  +-------- Rectidentini 
     |  |                      
     +--+     +-- Physunio **     
        |  +--+                
        |  |  +-- Trapezoideus ** 
        +--+                   
           |  +-- Yaukthwa **     
           +--+                
              +-- Lens **         

Konopleva et al., 2019 [Publ 3900], fig. 1

iTtTbiriSoTroTroSiD
16161443622217
  +----------- Rectidentini
--+                        
  |  +-------- Yaukthwa **    
  +--+                     
     |  +----- Physunio **    
     +--+                  
        |  +-- Trapezoideus **
        +--+               
           +-- Lens **        

Pfeiffer et al., 2021 [Publ 4126], fig. 9

iTtTbiriSoTroTroSiD
15151351414210
  +-------------- Lamprotulini
  |                           
--+  +----------- Rectidentini
  |  |                        
  +--+        +-- Trapezoideus **
     |  +-----+               
     |  |     +-- Physunio **    
     +--+                     
        |  +----- Yaukthwa **    
        +--+                  
           |  +-- Solenaia **    
           +--+               
              +-- Lens **        

Bolotov et al., 2022 [Publ 4250], fig. S5

iTtTbiriSoTroTroSiD
151512300014
  +----- Yaukthwa **    
--+                  
  |  +-- Physunio **    
  +--+               
     +-- Trapezoideus **

Zieritz et al., 2020 [Publ 4102], fig. 4

iTtTbiriSoTroTroSiD
14141241717615
  +----------------------- Chamberlainiini
  |                                       
--+  +-------------------- Lamprotulini   
  |  |                                    
  +--+  +----------------- Gonideini      
     |  |                                 
     +--+  +-------------- Pseudodontini  
        |  |                              
        +--+           +-- Ctenodesmini   
           |  +--------+                  
           |  |        +-- Rectidentini   
           +--+                           
              |  +-------- Physunio **       
              +--+                        
                 |  +----- Yaukthwa **       
                 +--+                     
                    |  +-- Trapezoideus **   
                    +--+                  
                       +-- Lens **           

Huang et al., 2019 [Publ 3843], fig. S4

iTtTbiriSoTroTroSiD
12127318744513
  +-------------- Rectidentini     
  |                                
  |  +----------- Physunio_modelli ** 
--+  |                             
  |  +----------- Lens **             
  |  |                             
  |  +----------- Physunio_superbus **
  +--+                             
     +----------- Yaukthwa **         
     |                             
     |        +-- Lens_comptus **     
     |  +-----+                    
     |  |     +-- Lens_novoselovi **  
     +--+                          
        |  +----- Lens_eximius **     
        +--+                       
           |  +-- Lens_contradens **  
           +--+                    
              +-- Lens_micropterus ** 

Bolotov et al., 2017 [Publ 3761], fig. S1

iTtTbiriSoTroTroSiD
121210312737512
  +-------- Rectidentini
--+                     
  |  +----- Physunio **    
  +--+                  
     |  +-- Yaukthwa **    
     +--+               
        +-- Lens **        

Huang et al., 2019 [Publ 3843], fig. 2

iTtTbiriSoTroTroSiD
121210318744512
  +-------- Rectidentini
--+                     
  |  +----- Physunio **    
  +--+                  
     |  +-- Yaukthwa **    
     +--+               
        +-- Lens **        

Bolotov et al., 2017 [Publ 3764], fig. S1

iTtTbiriSoTroTroSiD
121210311339512
  +-------- Rectidentini
--+                     
  |  +----- Physunio **    
  +--+                  
     |  +-- Yaukthwa **    
     +--+               
        +-- Lens **        

Zieritz et al., 2024 [Publ 4381], fig. 7A

iTtTbiriSoTroTroSiD
121210200013
  +-- Solenaia **
--+           
  +-- Lens **    

Huang et al., 2019 [Publ 3843], fig. S3

iTtTbiriSoTroTroSiD
121210318543512
  +-------- Rectidentini
--+                     
  |  +----- Physunio **    
  +--+                  
     |  +-- Yaukthwa **    
     +--+               
        +-- Lens **        

Bolotov et al., 2022 [Publ 4250], fig. 8

iTtTbiriSoTroTroSiD
12121023711313
  +----- Pseudodontini
--+                   
  |  +-- Trapezoideus ** 
  +--+                
     +-- Yaukthwa **     

Pfeiffer et al., 2021 [Publ 4126], fig. 12

iTtTbiriSoTroTroSiD
11118200012
  +-- Solenaia **
--+           
  +-- Lens **    

Bolotov et al., 2017 [Publ 3761], fig. S2

iTtTbiriSoTroTroSiD
11119312838513
  +-------- Rectidentini
--+                     
  |  +----- Lens **        
  +--+                  
     |  +-- Physunio **    
     +--+               
        +-- Yaukthwa **    

Jeratthitikul et al., 2021 [Publ 4212], fig. 4

iTtTbiriSoTroTroSiD
101084111118
  +-------- Rectidentini
  |                     
--+     +-- Trapezoideus **
  |  +--+               
  |  |  +-- Physunio **    
  +--+                  
     |  +-- Yaukthwa **    
     +--+               
        +-- Lens **        

Bolotov et al., 2018 [Publ 3826], fig. 2

iTtTbiriSoTroTroSiD
1010037325411
  +----- Rectidentini       
  |                         
  |  +-- Lens_eximius **       
  |  |                      
  |  +-- Physunio_modelli__1 **
  |  |                      
--+  +-- Yaukthwa_nesemanni ** 
  |  |                      
  |  +-- Yaukthwa_panhai **    
  |  |                      
  |  +-- Lens_comptus **       
  |  |                      
  +--+-- Lens_novoselovi **    
     |                      
     +-- Physunio_modelli__2 **
     |                      
     +-- Physunio **           
     |                      
     +-- Yaukthwa_inlenensis **
     |                      
     +-- Lens_contradens **    
     |                      
     +-- Lens_micropterus **   

Pfeiffer et al., 2021 [Publ 4126], fig. 25

iTtTbiriSoTroTroSiD
997200010
  +-- Solenaia **
--+           
  +-- Yaukthwa **

Pfeiffer et al., 2021 [Publ 4126], fig. 18

iTtTbiriSoTroTroSiD
88730008
  +----- Yaukthwa **    
--+                  
  |  +-- Trapezoideus **
  +--+               
     +-- Physunio **    

Zieritz et al., 2021 [Publ 4218], fig. 5

iTtTbiriSoTroTroSiD
7754252176
  +----------------------- Chamberlainiini
  |                                       
--+  +-------------------- Lamprotulini   
  |  |                                    
  |  |              +----- Gonideini      
  +--+  +-----------+                     
     |  |           |  +-- Schepmaniini   
     |  |           +--+                  
     +--+              +-- Pseudodontini  
        |                                 
        |  +-------------- Ctenodesmini   
        +--+                              
           |  +----------- Rectidentini   
           +--+                           
              |  +-------- Physunio **       
              +--+                        
                 |  +----- Trapezoideus **   
                 +--+                     
                    |  +-- Lens **           
                    +--+                  
                       +-- Yaukthwa **       

Liu et al., 2024 [Publ 4380], fig. 5

iTtTbiriSoTroTroSiD
7754302874
                    +-- Schepmaniini   
  +-----------------+                  
  |                 +-- Pseudodontini  
--+                                    
  |  +----------------- Gonideini      
  |  |                                 
  +--+              +-- Chamberlainiini
     |  +-----------+                  
     |  |           +-- Lamprotulini   
     +--+                              
        |  +----------- Ctenodesmini   
        |  |                           
        +--+  +-------- Rectidentini   
           |  |                        
           +--+     +-- Physunio **       
              |  +--+                  
              |  |  +-- Trapezoideus **   
              +--+                     
                 |  +-- Lens **           
                 +--+                  
                    +-- Yaukthwa **       

Wu et al., 2024 [Publ 4350], fig. 5c

iTtTbiriSoTroTroSiD
7714302876
  +----------------------------- Schepmania_nieuwenhuisi__1
  |                                                        
  +----------------------------- Schepmania_nieuwenhuisi__2
  |                                                        
  |  +-------------------------- Pseudodontini             
  |  |                                                     
--+  |                    +----- Ensidens                  
  |  |                    |                                
  |  |                    +----- Hyriopsis_desowitzi       
  |  |                    |                                
  |  |                    +----- Rectidens                 
  |  |                    |                                
  |  |                    +----- Hyriopsis_bialata         
  +--+  +-----------------+                                
     |  |                 +----- Ctenodesmini              
     |  |                 |                                
     |  |                 |  +-- Physunio **                  
     |  |                 |  |                             
     |  |                 |  +-- Yaukthwa_paiensis **         
     |  |                 |  |                             
     |  |                 |  +-- Yaukthwa_inlenensis **       
     |  |                 +--+                             
     +--+                    +-- Trapezoideus **              
        |                    |                             
        |                    +-- Yaukthwa_elongatula **       
        |                    |                             
        |                    +-- Lens **                      
        |                                                  
        |  +-------------------- Inversidens               
        |  |                                               
        |  |              +----- Lamprotula                
        +--+  +-----------+                                
           |  |           |  +-- Chamberlainiini           
           |  |           +--+                             
           +--+              +-- Potomida                  
              |                                            
              |  +-------------- Sinosolenaia              
              |  |                                         
              +--+  +----------- Gonidea                   
                 |  |                                      
                 +--+----------- Microcondylaea            
                    |                                      
                    |  +-------- Koreosolenaia             
                    +--+                                   
                       |  +----- Parvasolenaia             
                       +--+                                
                          |  +-- Postolata                 
                          +--+                             
                             +-- Ptychorhynchus            

Wu et al., 2024 [Publ 4350], fig. 6-7

iTtTbiriSoTroTroSiD
7754302874
  +----------------- Chamberlainiini
  |                                 
  |              +-- Schepmaniini   
  |           +--+                  
--+           |  +-- Pseudodontini  
  |  +--------+                     
  |  |        |  +-- Lamprotulini   
  |  |        +--+                  
  +--+           +-- Gonideini      
     |                              
     |  +----------- Ctenodesmini   
     |  |                           
     +--+  +-------- Rectidentini   
        |  |                        
        +--+     +-- Physunio **       
           |  +--+                  
           |  |  +-- Trapezoideus **   
           +--+                     
              |  +-- Lens **           
              +--+                  
                 +-- Yaukthwa **       

Wu et al., 2024 [Publ 4350], fig. 5a

iTtTbiriSoTroTroSiD
7754302874
  +-------------------------- Chamberlainiini
  |                                          
  |              +----------- Ctenodesmini   
  |              |                           
  |  +-----------+  +-------- Rectidentini   
--+  |           |  |                        
  |  |           +--+     +-- Physunio **       
  |  |              |  +--+                  
  |  |              |  |  +-- Trapezoideus **   
  |  |              +--+                     
  |  |                 |  +-- Lens **           
  +--+                 +--+                  
     |                    +-- Yaukthwa **       
     |                                       
     |                    +-- Schepmaniini   
     |  +-----------------+                  
     |  |                 +-- Pseudodontini  
     |  |                                    
     +--+                 +-- Inversidens    
        |  +--------------+                  
        |  |              +-- Lamprotulini   
        +--+                                 
           |  +-------------- Sinosolenaia   
           |  |                              
           +--+           +-- Gonidea        
              |  +--------+                  
              |  |        +-- Microcondylaea 
              +--+                           
                 |  +-------- Parvasolenaia  
                 +--+                        
                    |  +----- Koreosolenaia  
                    +--+                     
                       |  +-- Postolata      
                       +--+                  
                          +-- Ptychorhynchus 

Wu et al., 2024 [Publ 4388], fig. 5

iTtTbiriSoTroTroSiD
7754312974
  +----------------- Chamberlainiini
  |                                 
  |        +-------- Lamprotulini   
--+  +-----+                        
  |  |     |  +----- Gonideini      
  |  |     +--+                     
  |  |        |  +-- Schepmaniini   
  +--+        +--+                  
     |           +-- Pseudodontini  
     |                              
     |  +----------- Ctenodesmini   
     |  |                           
     +--+  +-------- Rectidentini   
        |  |                        
        +--+     +-- Physunio **       
           |  +--+                  
           |  |  +-- Trapezoideus **   
           +--+                     
              |  +-- Lens **           
              +--+                  
                 +-- Yaukthwa **       

Wu et al., 2024 [Publ 4350], fig. 5b

iTtTbiriSoTroTroSiD
7754302876
  +-------------------------------- Schepmaniini       
  |                                                    
  |  +----------------------------- Pseudodontini      
  |  |                                                 
--+  |     +----------------------- Hyriopsis_bialata  
  |  |     |                                           
  |  |  +--+                    +-- Rectidens          
  |  |  |  |  +-----------------+                      
  |  |  |  |  |                 +-- Ctenodesmini       
  +--+  |  +--+                                        
     |  |     |  +----------------- Ensidens           
     |  |     |  |                                     
     |  |     +--+  +-------------- Hyriopsis_desowitzi
     |  |        |  |                                  
     |  |        +--+  +----------- Yaukthwa_elongatula **
     |  |           |  |                               
     |  |           +--+  +-------- Trapezoideus **       
     +--+              |  |                            
        |              +--+     +-- Physunio **           
        |                 |  +--+                      
        |                 |  |  +-- Yaukthwa_inlenensis **
        |                 +--+                         
        |                    |  +-- Yaukthwa_paiensis **  
        |                    +--+                      
        |                       +-- Lens **               
        |                                              
        |     +-------------------- Inversidens        
        |     |                                        
        |     |              +----- Lamprotula         
        +-----+  +-----------+                         
              |  |           |  +-- Chamberlainiini    
              |  |           +--+                      
              +--+              +-- Potomida           
                 |                                     
                 |  +-------------- Sinosolenaia       
                 |  |                                  
                 +--+           +-- Gonidea            
                    |  +--------+                      
                    |  |        +-- Microcondylaea     
                    +--+                               
                       |  +-------- Koreosolenaia      
                       +--+                            
                          |  +----- Parvasolenaia      
                          +--+                         
                             |  +-- Postolata          
                             +--+                      
                                +-- Ptychorhynchus     

Jeratthitikul et al., 2019 [Publ 3911], fig. 4

iTtTbiriSoTroTroSiD
6611232218
  +----- Rectidentini
--+                  
  |  +-- Physunio **    
  +--+               
     +-- Lens **        

Pfeiffer et al., 2019 [Publ 3937], fig. 6

iTtTbiriSoTroTroSiD
5535521250
  +----------------- Chamberlainiini
  |                                 
  |           +----- Pseudodontini  
--+  +--------+                     
  |  |        |  +-- Gonideini      
  |  |        +--+                  
  +--+           +-- Lamprotulini   
     |                              
     |  +----------- Rectidentini   
     |  |                           
     +--+        +-- Trapezoideus **   
        |  +-----+                  
        |  |     +-- Physunio **       
        +--+                        
           |  +----- Yaukthwa **       
           +--+                     
              |  +-- Lens **           
              +--+                  
                 +-- Solenaia **       

Bolotov et al., 2019 [Publ 3974], fig. 1

iTtTbiriSoTroTroSiD
553113009
1

Wu et al., 2021 [Publ 4366], fig. 2_I

iTtTbiriSoTroTroSiD
4423872556
  +-------- Rectidentini
--+                     
  |  +----- Physunio **    
  +--+                  
     |  +-- Lens **        
     +--+               
        +-- Yaukthwa **    

Wu et al., 2021 [Publ 4366], fig. 6

iTtTbiriSoTroTroSiD
4423872556
  +-------- Rectidentini
--+                     
  |  +----- Physunio **    
  +--+                  
     |  +-- Lens **        
     +--+               
        +-- Yaukthwa **    

Pfeiffer et al., 2018 [Publ 3827], fig. 1

iTtTbiriSoTroTroSiD
44133315
  +----- Rectidentini
  |                  
--+  +-- Physunio **    
  |  |               
  +--+-- Trapezoideus **
     |               
     +-- Lens **        

Jeratthitikul et al., 2024 [Publ 4387], fig. 1

iTtTbiriSoTroTroSiD
3313555574
           +-------- Lamprotulini   
  +--------+                        
  |        |  +----- Gonideini      
  |        +--+                     
  |           |  +-- Schepmaniini   
--+           +--+                  
  |              +-- Pseudodontini  
  |                                 
  |  +-------------- Chamberlainiini
  +--+                              
     |  +----------- Ctenodesmini   
     +--+                           
        |  +-------- Rectidentini   
        +--+                        
           |  +----- Physunio **       
           +--+                     
              |  +-- Lens **           
              +--+                  
                 +-- Trapezoideus **   

Konopleva et al., 2017 [Publ 3807], fig. 6

iTtTbiriSoTroTroSiD
331228725.5
  +-------- Pseudodontini
--+                      
  |  +----- Rectidentini 
  +--+                   
     |  +-- Lens **         
     +--+                
        +-- Physunio **     

Lopes-Lima et al., 2017 [Publ 3745], fig. 1-2

iTtTbiriSoTroTroSiD
3422772456
  +-------- Rectidentini
--+                     
  |  +----- Lens **        
  +--+                  
     |  +-- Physunio **    
     +--+               
        +-- unionoid    

Dai et al., 2024 [Publ 4389], fig. 3

iTtTbiriSoTroTroSiD
3312443974
        +-------- Ctenodesmini   
  +-----+                        
  |     |  +----- Rectidentini   
  |     +--+                     
  |        |  +-- Physunio **       
--+        +--+                  
  |           +-- Lens **           
  |                              
  |  +----------- Chamberlainiini
  +--+                           
     |  +-------- Lamprotulini   
     +--+                        
        |  +----- Gonideini      
        +--+                     
           |  +-- Schepmaniini   
           +--+                  
              +-- Pseudodontini  

Jeratthitikul et al., 2021 [Publ 4219], fig. 2

iTtTbiriSoTroTroSiD
3313333352
     +-------- Rectidentini   
  +--+                        
  |  |  +----- Lens **           
  |  +--+                     
  |     |  +-- Trapezoideus **   
--+     +--+                  
  |        +-- Physunio **       
  |                           
  |  +-------- Pseudodontini  
  +--+                        
     |  +----- Gonideini      
     +--+                     
        |  +-- Chamberlainiini
        +--+                  
           +-- Lamprotulini   

Dai et al., 2023 [Publ 4285], fig. 3-ML

iTtTbiriSoTroTroSiD
3312424054
  +----------------------- Pseudodontini  
  |                                       
  |                 +----- Rectidentini   
  |  +--------------+                     
--+  |              |  +-- Physunio **       
  |  |              +--+                  
  |  |                 +-- Lens **           
  |  |                                    
  +--+              +----- Inversidens    
     |  +-----------+                     
     |  |           |  +-- Chamberlainiini
     |  |           +--+                  
     |  |              +-- Lamprotulini   
     +--+                                 
        |  +-------------- Sinosolenaia   
        |  |                              
        |  |        +----- Gonidea        
        +--+  +-----+                     
           |  |     |  +-- Leguminaia     
           |  |     +--+                  
           |  |        +-- Microcondylaea 
           +--+                           
              |        +-- Koreosolenaia  
              |  +-----+                  
              |  |     +-- Parvasolenaia  
              +--+                        
                 |  +----- Postolata      
                 +--+                     
                    |  +-- Ptychorhynchus 
                    +--+                  
                       +-- Obovalis       

Jeratthitikul et al., 2021 [Publ 4219], fig. 1

iTtTbiriSoTroTroSiD
3313333352
     +-------- Rectidentini   
  +--+                        
  |  |  +----- Lens **           
  |  +--+                     
  |     |  +-- Trapezoideus **   
--+     +--+                  
  |        +-- Physunio **       
  |                           
  |  +-------- Pseudodontini  
  +--+                        
     |  +----- Gonideini      
     +--+                     
        |  +-- Chamberlainiini
        +--+                  
           +-- Lamprotulini   

Pfeiffer & Graf, 2015 [Publ 3620], fig. 2

iTtTbiriSoTroTroSiD
3312371145.5
     +----- Rectidentini 
  +--+                   
  |  |  +-- Lens **         
  |  +--+                
--+     +-- Physunio **     
  |                      
  |  +----- Pseudodontini
  +--+                   
     |  +-- Gonideini    
     +--+                
        +-- Lamprotulini 

Dai et al., 2025 [Publ 4441], fig. 1

iTtTbiriSoTroTroSiD
3312454074
        +-------- Ctenodesmini   
  +-----+                        
  |     |  +----- Rectidentini   
  |     +--+                     
  |        |  +-- Physunio **       
--+        +--+                  
  |           +-- Lens **           
  |                              
  |  +----------- Chamberlainiini
  +--+                           
     |  +-------- Lamprotulini   
     +--+                        
        |  +----- Gonideini      
        +--+                     
           |  +-- Schepmaniini   
           +--+                  
              +-- Pseudodontini  

Lopes-Lima et al., 2020 [Publ 4080], fig. 2-4

iTtTbiriSoTroTroSiD
35031183057
  +-- Lens **     
  |            
  +-- Ensidens 
  |            
--+-- Physunio ** 
  |            
  +-- Rectidens
  |            
  +-- Yaukthwa ** 

Jeratthitikul et al., 2022 [Publ 4260], fig. S1_1

iTtTbiriSoTroTroSiD
3313732852
  +-------- Rectidentini
--+                     
  |  +----- Lens **        
  +--+                  
     |  +-- Trapezoideus **
     +--+               
        +-- Physunio **    

Jeratthitikul et al., 2022 [Publ 4260], fig. S2_2

iTtTbiriSoTroTroSiD
3313732852
  +-------- Rectidentini
--+                     
  |  +----- Lens **        
  +--+                  
     |  +-- Trapezoideus **
     +--+               
        +-- Physunio **    

Dai et al., 2023 [Publ 4285], fig. 3-BI

iTtTbiriSoTroTroSiD
3312424054
  +----------------------- Pseudodontini  
  |                                       
  |                 +----- Rectidentini   
--+  +--------------+                     
  |  |              |  +-- Physunio **       
  |  |              +--+                  
  |  |                 +-- Lens **           
  +--+                                    
     |  +----------------- Inversidens    
     |  |                                 
     |  |              +-- Chamberlainiini
     |  +--------------+                  
     +--+              +-- Lamprotulini   
        |                                 
        |  +-------------- Sinosolenaia   
        |  |                              
        |  |        +----- Gonidea        
        +--+  +-----+                     
           |  |     |  +-- Leguminaia     
           |  |     +--+                  
           |  |        +-- Microcondylaea 
           +--+                           
              |        +-- Koreosolenaia  
              |  +-----+                  
              |  |     +-- Parvasolenaia  
              +--+                        
                 |  +----- Postolata      
                 +--+                     
                    |  +-- Ptychorhynchus 
                    +--+                  
                       +-- Obovalis       

Zieritz et al., 2021 [Publ 4231], fig. 3-M_4

iTtTbiriSoTroTroSiD
220221105--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Pseudodontini  
     +--+                     
        |  +-- Lamprotulini   
        +--+                  
           +-- Gonideini      

Dai et al., 2025 [Publ 4440], fig. 3

iTtTbiriSoTroTroSiD
220286175--
  +----- Rectidentini
--+                  
  |  +-- Lens **        
  +--+               
     +-- Physunio **    

Zieritz et al., 2021 [Publ 4231], fig. 3-F_5

iTtTbiriSoTroTroSiD
220246155--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Wu et al., 2024 [Publ 4388], fig. 6

iTtTbiriSoTroTroSiD
220252215--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Wu et al., 2023 [Publ 4339], fig. 4_S5-aa

iTtTbiriSoTroTroSiD
220293185--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Wu et al., 2024 [Publ 4350], fig. 8b_S2

iTtTbiriSoTroTroSiD
220289195--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Dai et al., 2023 [Publ 4285], fig. 5

iTtTbiriSoTroTroSiD
220251175--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Wu et al., 2024 [Publ 4350], fig. 8a_S1

iTtTbiriSoTroTroSiD
220289195--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Matos et al., 2024 [Publ 4384], fig. 3

iTtTbiriSoTroTroSiD
220218165--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Physunio **       
  |  |  +--+                  
  +--+     +-- Lens **           
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Zhang et al., 2024 [Publ 4397], fig. 5

iTtTbiriSoTroTroSiD
220226245--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Wu et al., 2023 [Publ 4339], fig. 4_S5-nt

iTtTbiriSoTroTroSiD
220293185--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Lens **           
  |  |  +--+                  
  +--+     +-- Physunio **       
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Matos et al., 2024 [Publ 4383], fig. 3

iTtTbiriSoTroTroSiD
220218165--
  +----------- Chamberlainiini
  |                           
  |     +----- Rectidentini   
--+  +--+                     
  |  |  |  +-- Physunio **       
  |  |  +--+                  
  +--+     +-- Lens **           
     |                        
     |  +----- Lamprotulini   
     +--+                     
        |  +-- Pseudodontini  
        +--+                  
           +-- Gonideini      

Pfeiffer & Graf, 2013 [Publ 3531], fig. 2

iTtTbiriSoTroTroSiD
110130104--
  +-------- Chamberlainiini
  |                        
--+     +-- Gonideini      
  |  +--+                  
  |  |  +-- Pseudodontini  
  +--+                     
     |  +-- Physunio **       
     +--+                  
        +-- Lamprotulini   

Hoeh et al., 2009 [Publ 2716], fig. 4

iTtTbiriSoTroTroSiD
11014532--
-- Lens_contradens **

Graf & Cummings, 2006 [Publ 1968], fig. 2

iTtTbiriSoTroTroSiD
11014632--
-- Lens_contradens **

Whelan et al., 2011 [Publ 3380], fig. 1-ML

iTtTbiriSoTroTroSiD
11013553--
  +-- Pseudodontini
--+                
  +-- Lens **         

Hoeh et al., 2009 [Publ 2716], fig. 5

iTtTbiriSoTroTroSiD
11014532--
-- Lens_contradens **

Hoeh et al., 2009 [Publ 2716], fig. 3

iTtTbiriSoTroTroSiD
11014632--
-- Lens_contradens **

Whelan et al., 2011 [Publ 3380], fig. 1-MP

iTtTbiriSoTroTroSiD
11013553--
-- Lens_contradens **

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
11011922--
  +----- Chamberlainiini
--+                     
  |  +-- Lens **           
  +--+                  
     +-- Lamprotulini   

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
11011732--
-- Lens_contradens **
 
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