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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 | Contradentini

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

Cladomics Summary

number of species = 4 (4 total)
analyzed in 57 cladograms in 38 publications
maximum number of terminals in an analysis = 4 (mean = 1.6, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 20
maximum number of included terminals = 4, (mean = 2.7 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 **

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 **

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

iTtTbiriSoTroTroSiD
3313196925
        +-- 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 **

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

iTtTbiriSoTroTroSiD
3313791633
           +-- Yaukthwa         
  +--------+                    
  |        +-- Lens             
--+                             
  |  +-------- Trapezoideus     
  +--+                          
     |  +----- Physunio_modelli ** 
     +--+                       
        |  +-- Physunio_superbus **
        +--+                    
           +-- Physunio_massini **

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

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

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 **

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

iTtTbiriSoTroTroSiD
3313194925
        +-- Yaukthwa         
  +-----+                    
  |     +-- Lens             
--+                          
  |  +----- Physunio **         
  +--+                       
     |  +-- Physunio_superbus **
     +--+                    
        +-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
2120219710213
  +----------- Physunio_modelli ** 
  |                             
  +----------- Lens             
  |                             
  +----------- Physunio_superbus **
--+                             
  +----------- Yaukthwa         
  |                             
  |        +-- Lens_comptus     
  |  +-----+                    
  |  |     +-- Lens_novoselovi  
  +--+                          
     |  +----- Lens_eximius     
     +--+                       
        |  +-- Lens_contradens  
        +--+                    
           +-- Lens_micropterus

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
11013000--
-- Physunio **

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

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

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

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

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

iTtTbiriSoTroTroSiD
11015722--
  +----- Physunio_superbus **
--+                       
  |  +-- Lens             
  +--+                    
     +-- Trapezoideus

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

iTtTbiriSoTroTroSiD
11017921--
-- Physunio_modelli **

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

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

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

iTtTbiriSoTroTroSiD
11015311--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

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

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

iTtTbiriSoTroTroSiD
11019411--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

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

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

iTtTbiriSoTroTroSiD
11019011--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
11019011--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

iTtTbiriSoTroTroSiD
11012711--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

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

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

iTtTbiriSoTroTroSiD
11019411--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

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

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

iTtTbiriSoTroTroSiD
110112022--
-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
11013763--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

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

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

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

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

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

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

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

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

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
 
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