MUSSELp
 
MUSSELpdb
  Introduction
Browse
d'basing
 
Taxonomy
  all genera
Unionidae
Margaritiferidae
Hyriidae
Etheriidae
Mycetopodidae
Iridinidae

Species & Genus Richness by Taxonomy
 
Geography
  Summary
North America
Central America
South America
Afrotropics
North Eurasia
East Asia
Australasia

Species Richness & Endemism by Geography
 
 

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 65 cladograms in 43 publications
maximum number of terminals in an analysis = 5 (mean = 1.62, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 22
maximum number of included terminals = 5, (mean = 2.82 median = 3)
proportion of cladograms that support basic monophyly = 0.86

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 **

Campbell & Lydeard, 2026 [Publ 4810], fig. 6-14

iTtTbiriSoTroTroSiD
53130562726329
              +-- Physunio_modelli **   
  +-----------+                      
  |           +-- Physunio_pellucidus **
  |                                  
--+           +-- Physunio_massini **   
  |  +--------+                      
  |  |        +-- Physunio_superbus **  
  +--+                               
     |  +-------- Physunio **           
     +--+                            
        |  +----- Trapezoideus       
        +--+                         
           |  +-- Yaukthwa           
           +--+                      
              +-- Lens

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

iTtTbiriSoTroTroSiD
3313135825
  +----------- Lens             
--+                             
  |  +-------- Yaukthwa         
  +--+                          
     |  +----- Physunio **         
     +--+                       
        |  +-- Physunio_modelli ** 
        +--+                    
           +-- Physunio_superbus **

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

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

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

iTtTbiriSoTroTroSiD
312731979213
  +-- Physunio_modelli ** 
  |                    
  +-- Physunio **         
  |                    
--+-- Physunio_superbus **
  |                    
  +-- Yaukthwa         
  |                    
  +-- Lens

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

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

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

iTtTbiriSoTroTroSiD
33131082035
     +----- Physunio **         
  +--+                       
  |  |  +-- Physunio_modelli ** 
  |  +--+                    
--+     +-- Physunio_superbus **
  |                          
  |  +----- Trapezoideus     
  +--+                       
     |  +-- Lens             
     +--+                    
        +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
3313135925
        +-- 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
3313195925
        +-- Yaukthwa         
  +-----+                    
  |     +-- Lens             
--+                          
  |  +----- Physunio **         
  +--+                       
     |  +-- Physunio_superbus **
     +--+                    
        +-- Physunio_modelli **

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

iTtTbiriSoTroTroSiD
3313197925
        +-- Yaukthwa         
  +-----+                    
  |     +-- Lens             
--+                          
  |  +----- Physunio **         
  +--+                       
     |  +-- Physunio_modelli ** 
     +--+                    
        +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
3313481335
  +----------- Yaukthwa         
  |                             
--+        +-- Trapezoideus     
  |  +-----+                    
  |  |     +-- Lens             
  +--+                          
     |  +----- Physunio_superbus **
     +--+                       
        |  +-- Physunio_modelli ** 
        +--+                    
           +-- Physunio **

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

iTtTbiriSoTroTroSiD
33131142035
           +-- Yaukthwa         
  +--------+                    
  |        +-- Lens             
--+                             
  |  +-------- Trapezoideus     
  +--+                          
     |  +----- Physunio_superbus **
     +--+                       
        |  +-- Physunio_modelli ** 
        +--+                    
           +-- Physunio **

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

iTtTbiriSoTroTroSiD
3313121925
        +-- Yaukthwa         
  +-----+                    
  |     +-- Lens             
--+                          
  |  +----- Physunio **         
  +--+                       
     |  +-- Physunio_superbus **
     +--+                    
        +-- Physunio_modelli **

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
21003808213
  +-- Lens_eximius       
  |                      
  +-- Physunio_modelli__1 **
  |                      
  +-- Yaukthwa_nesemanni 
  |                      
  +-- Yaukthwa_panhai    
  |                      
  +-- Lens_comptus       
  |                      
--+-- Lens_novoselovi    
  |                      
  +-- Physunio_modelli__2 **
  |                      
  +-- Physunio **           
  |                      
  +-- Yaukthwa_inlenensis
  |                      
  +-- Lens_contradens    
  |                      
  +-- Lens_micropterus

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
11013563--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

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

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

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

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

Wu et al., 2026 [Publ 4811], fig. 4

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
11015822--
  +----- Physunio_superbus **
--+                       
  |  +-- Lens             
  +--+                    
     +-- Trapezoideus

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

iTtTbiriSoTroTroSiD
11013563--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
11013563--
  +----------- Yaukthwa_elongatula
  |                               
--+  +-------- Trapezoideus       
  |  |                            
  +--+     +-- Physunio_superbus **  
     |  +--+                      
     |  |  +-- Yaukthwa_inlenensis
     +--+                         
        |  +-- Yaukthwa_paiensis  
        +--+                      
           +-- Lens

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

iTtTbiriSoTroTroSiD
11013063--
  +-------- Physunio_superbus **
--+                          
  |  +----- Trapezoideus     
  +--+                       
     |  +-- Lens             
     +--+                    
        +-- Yaukthwa

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

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

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

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

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

iTtTbiriSoTroTroSiD
11014473--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Yaukthwa         
  +--+                    
     +-- Lens

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

iTtTbiriSoTroTroSiD
11014521--
  +-- Physunio_superbus **
--+                    
  +-- Lens

Klymus et al., 2025 [Publ 4814], fig. 3B

iTtTbiriSoTroTroSiD
11013511--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

iTtTbiriSoTroTroSiD
11013322--
  +----- Lens             
--+                       
  |  +-- Trapezoideus     
  +--+                    
     +-- Physunio_superbus **

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
11013000--
-- Physunio **

Campbell & Lydeard, 2026 [Publ 4810], fig. 4

iTtTbiriSoTroTroSiD
11014611--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

iTtTbiriSoTroTroSiD
11013563--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

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

iTtTbiriSoTroTroSiD
11013563--
     +-- Physunio_superbus **
  +--+                    
  |  +-- Trapezoideus     
--+                       
  |  +-- Lens             
  +--+                    
     +-- Yaukthwa

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

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

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

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

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

iTtTbiriSoTroTroSiD
110112122--
-- Physunio_modelli **

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

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

Campbell & Lydeard, 2026 [Publ 4810], fig. 3_5

iTtTbiriSoTroTroSiD
110114011--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

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

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

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

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

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

Klymus et al., 2025 [Publ 4814], fig. 3A

iTtTbiriSoTroTroSiD
11013511--
  +-- Lens             
--+                    
  +-- Physunio_superbus **

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
11013563--
  +-- Physunio_superbus **  
  |                      
  +-- Yaukthwa_paiensis  
  |                      
  +-- Yaukthwa_inlenensis
--+                      
  +-- Trapezoideus       
  |                      
  +-- Yaukthwa_elongatula
  |                      
  +-- Lens

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

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

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

iTtTbiriSoTroTroSiD
11019111--
  +-- Lens             
--+                    
  +-- Physunio_superbus **
 
NSF icon MUSSEL icon
"Making the world a better place, one mollusk at a time."