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Page last updated
Thu 3 September 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!

Search the database pages:

Bivalvia | Unionida | Unionoidea | Unionidae | Gonideinae | Rectidentini | Contradentina

genus Physunio Simpson, 1900: cladomics
     4 Recent valid species (4 total species)

Cladomics Summary

number of species = 4 (4 total)
analyzed in 59 cladograms in 40 publications
maximum number of terminals in an analysis = 4 (mean = 1.61, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 21
maximum number of included terminals = 4, (mean = 2.71 median = 3)
proportion of cladograms that support basic monophyly = 0.9

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., 2021 [Publ 4126], fig. 18

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
44364420
  +-------------- Yaukthwa           
  |                                  
--+  +----------- Trapezoideus       
  |  |                               
  +--+        +-- Physunio_massini **   
     |  +-----+                      
     |  |     +-- Physunio_superbus **  
     +--+                            
        |  +----- Physunio_modelli__3 **
        |  |                         
        +--+  +-- Physunio_modelli__1 **
           |  |                      
           +--+-- Physunio_modelli__2 **
              |                      
              +-- Physunio_pellucidus **

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

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
3313261243
        +----- Yaukthwa           
  +-----+                         
  |     |  +-- Solenaia           
  |     +--+                      
--+        +-- Lens               
  |                               
  |  +-------- Trapezoideus       
  +--+                            
     |  +----- Physunio_superbus **  
     +--+                         
        |  +-- Physunio_modelli **   
        +--+                      
           +-- Physunio_pellucidus **

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

iTtTbiriSoTroTroSiD
3313198925
        +-- Yaukthwa         
  +-----+                    
  |     +-- Lens             
--+                          
  |  +----- Physunio **         
  +--+                       
     |  +-- Physunio_modelli ** 
     +--+                    
        +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
331318733
           +-- Yaukthwa           
  +--------+                      
  |        +-- Lens               
--+                               
  |  +-------- Trapezoideus       
  +--+                            
     |  +----- Physunio_superbus **  
     +--+                         
        |  +-- Physunio_pellucidus **
        +--+                      
           +-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
3313122925
        +-- Yaukthwa         
  +-----+                    
  |     +-- Lens             
--+                          
  |  +----- Physunio **         
  +--+                       
     |  +-- Physunio_superbus **
     +--+                    
        +-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
3313136825
  +----------- Lens             
--+                             
  |  +-------- Yaukthwa         
  +--+                          
     |  +----- Physunio **         
     +--+                       
        |  +-- Physunio_modelli ** 
        +--+                    
           +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
3313801633
           +-- Yaukthwa         
  +--------+                    
  |        +-- Lens             
--+                             
  |  +-------- Trapezoideus     
  +--+                          
     |  +----- Physunio_modelli ** 
     +--+                       
        |  +-- Physunio_superbus **
        +--+                    
           +-- Physunio_massini **

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

iTtTbiriSoTroTroSiD
312731989213
  +-- Physunio_modelli ** 
  |                    
  +-- Physunio **         
  |                    
--+-- Physunio_superbus **
  |                    
  +-- Yaukthwa         
  |                    
  +-- Lens

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

iTtTbiriSoTroTroSiD
33131092035
     +----- Physunio **         
  +--+                       
  |  |  +-- Physunio_modelli ** 
  |  +--+                    
--+     +-- Physunio_superbus **
  |                          
  |  +----- Trapezoideus     
  +--+                       
     |  +-- Lens             
     +--+                    
        +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
3313136925
        +-- Yaukthwa         
  +-----+                    
  |     +-- Lens             
--+                          
  |  +----- Physunio **         
  +--+                       
     |  +-- Physunio_modelli ** 
     +--+                    
        +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
3313311635
  +-------------- Yaukthwa         
--+                                
  |  +----------- Lens             
  +--+                             
     |  +-------- Trapezoideus     
     +--+                          
        |  +----- Physunio_superbus **
        +--+                       
           |  +-- Physunio_modelli ** 
           +--+                    
              +-- Physunio **

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

iTtTbiriSoTroTroSiD
3313196925
        +-- Yaukthwa         
  +-----+                    
  |     +-- Lens             
--+                          
  |  +----- Physunio **         
  +--+                       
     |  +-- Physunio_superbus **
     +--+                    
        +-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
3313491335
  +----------- Yaukthwa         
  |                             
--+        +-- Trapezoideus     
  |  +-----+                    
  |  |     +-- Lens             
  +--+                          
     |  +----- Physunio_superbus **
     +--+                       
        |  +-- Physunio_modelli ** 
        +--+                    
           +-- Physunio **

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

iTtTbiriSoTroTroSiD
33131152035
           +-- Yaukthwa         
  +--------+                    
  |        +-- Lens             
--+                             
  |  +-------- Trapezoideus     
  +--+                          
     |  +----- Physunio_superbus **
     +--+                       
        |  +-- Physunio_modelli ** 
        +--+                    
           +-- Physunio **

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

iTtTbiriSoTroTroSiD
220229123--
        +-- Physunio_modelli ** 
  +-----+                    
  |     +-- Physunio_superbus **
--+                          
  |  +----- Yaukthwa         
  +--+                       
     |  +-- Trapezoideus     
     +--+                    
        +-- Lens

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

iTtTbiriSoTroTroSiD
22023811--
  +----- Lens            
--+                      
  |  +-- Physunio **        
  +--+                   
     +-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
22027821--
  +----- Lens            
--+                      
  |  +-- Physunio_modelli **
  +--+                   
     +-- Physunio **

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

iTtTbiriSoTroTroSiD
22022911--
  +----- Lens            
--+                      
  |  +-- Physunio **        
  +--+                   
     +-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
2202741--
  +----- Lens             
--+                       
  |  +-- Physunio_modelli ** 
  +--+                    
     +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
220213132--
  +-------- Yaukthwa         
--+                          
  |  +----- Trapezoideus     
  +--+                       
     |  +-- Physunio_modelli ** 
     +--+                    
        +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
21003818213
  +-- Lens_eximius       
  |                      
  +-- Physunio_modelli__1 **
  |                      
  +-- Yaukthwa_nesemanni 
  |                      
  +-- Yaukthwa_panhai    
  |                      
  +-- Lens_comptus       
  |                      
--+-- Lens_novoselovi    
  |                      
  +-- Physunio_modelli__2 **
  |                      
  +-- Physunio **           
  |                      
  +-- Yaukthwa_inlenensis
  |                      
  +-- Lens_contradens    
  |                      
  +-- Lens_micropterus

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

iTtTbiriSoTroTroSiD
11017522--
  +----- Lens             
--+                       
  |  +-- Trapezoideus     
  +--+                    
     +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11013163--
  +-------- Physunio_superbus **
--+                          
  |  +----- Trapezoideus     
  +--+                       
     |  +-- Lens             
     +--+                    
        +-- Yaukthwa

Chen et al., 2025 [Publ 4663], fig. 1

iTtTbiriSoTroTroSiD
11014821--
  +-- Physunio_superbus **
--+                    
  +-- Lens

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

iTtTbiriSoTroTroSiD
11019032--
  +----- Physunio_modelli **
--+                      
  |  +-- Lens            
  +--+                   
     +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
11013663--
  +----------- Yaukthwa_elongatula
  |                               
--+  +-------- Trapezoideus       
  |  |                            
  +--+     +-- Physunio_superbus **  
     |  +--+                      
     |  |  +-- Yaukthwa_inlenensis
     +--+                         
        |  +-- Yaukthwa_paiensis  
        +--+                      
           +-- Lens

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

iTtTbiriSoTroTroSiD
11013663--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
11014421--
  +-- Physunio_superbus **
--+                    
  +-- Lens

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

iTtTbiriSoTroTroSiD
11014711--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11013522--
  +----- Lens             
--+                       
  |  +-- Trapezoideus     
  +--+                    
     +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11019511--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11011911--
  +-- Physunio_superbus **
--+                    
  +-- Lens

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

iTtTbiriSoTroTroSiD
11013663--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
11012211--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11019111--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11015922--
  +----- Physunio_superbus **
--+                       
  |  +-- Lens             
  +--+                    
     +-- Trapezoideus

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

iTtTbiriSoTroTroSiD
11018711--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11019511--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11012611--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11013663--
  +-- Physunio_superbus **  
  |                      
  +-- Yaukthwa_paiensis  
  |                      
  +-- Yaukthwa_inlenensis
--+                      
  +-- Trapezoideus       
  |                      
  +-- Yaukthwa_elongatula
  |                      
  +-- Lens

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

iTtTbiriSoTroTroSiD
110112122--
-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
11015644--
        +-- Trapezoideus     
  +-----+                    
  |     +-- Physunio_superbus **
--+                          
  |  +----- Yaukthwa         
  +--+                       
     |  +-- Lens             
     +--+                    
        +-- Solenaia

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

iTtTbiriSoTroTroSiD
11014621--
  +-- Physunio_superbus **
--+                    
  +-- Lens

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

iTtTbiriSoTroTroSiD
11013422--
  +----- Lens             
--+                       
  |  +-- Trapezoideus     
  +--+                    
     +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11019032--
  +----- Physunio_modelli **
--+                      
  |  +-- Lens            
  +--+                   
     +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
11017522--
  +----- Lens             
--+                       
  |  +-- Trapezoideus     
  +--+                    
     +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11013663--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
11015644--
        +-- Trapezoideus     
  +-----+                    
  |     +-- Physunio_superbus **
--+                          
  |  +----- Yaukthwa         
  +--+                       
     |  +-- Lens             
     +--+                    
        +-- Solenaia

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

iTtTbiriSoTroTroSiD
11014421--
  +-- Physunio_superbus **
--+                    
  +-- Lens

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

iTtTbiriSoTroTroSiD
11014573--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Yaukthwa         
  +--+                    
     +-- Lens

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11014721--
  +-- Physunio_superbus **
--+                    
  +-- Lens

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

iTtTbiriSoTroTroSiD
1101632--
  +-- Physunio **    
  |               
--+-- Trapezoideus
  |               
  +-- Lens

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

iTtTbiriSoTroTroSiD
11019111--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11013663--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

Chen et al., 2025 [Publ 4448], fig. 3

iTtTbiriSoTroTroSiD
11018511--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11011911--
  +-- Physunio_superbus **
--+                    
  +-- Lens

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

iTtTbiriSoTroTroSiD
11013000--
-- Physunio **
 
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