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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!

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Bivalvia | Unionida | Hyrioidea | Hyriidae

subfamily VELESUNIONINAE Iredale, 1934: cladomics
     6 genera

Cladomics Summary

number of species = 17 (21 total)
number of subtaxa (rank: genus) = 6
analyzed in 65 cladograms in 38 publications
maximum number of terminals in an analysis = 10 (mean = 2.38, median = 1)
maximum number of subtaxa (genus) = 5 (mean = 1.86, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 27
maximum number of included terminals = 10, (mean = 4.33 median = 3)
maximum number of subtaxa (genus) = 5 (mean = 3.07, median = 2)
proportion of cladograms that support basic monophyly = 0.67

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 26
(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 = 10 (mean = 4.42, median = 3)
maximum number of subtaxa (genus) = 5 (mean = 3.15, median = 2)
proportion of cladograms that support rigorous monophyly = 0.65


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


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

** Top Monophyly Cladogram ** ** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
10283456221810
  +----------------------------------- Westralunio **           
  |                                                          
  |                             +----- Diplodon_suavidicus__2
  |  +--------------------------+                            
--+  |                          |  +-- Hyriini               
  |  |                          +--+                         
  |  |                             +-- Castaliini            
  |  |                                                       
  |  |                             +-- Hyridella_depressa    
  +--+                       +-----+                         
     |                       |     +-- Hyridella_drapeta     
     |  +--------------------+                               
     |  |                    |  +----- Diplodon_suavidicus__1
     |  |                    +--+                            
     |  |                       |  +-- Diplodon_chilensis    
     |  |                       +--+                         
     +--+                          +-- Diplodon_gratus       
        |                                                    
        |                          +-- Cucumerunio           
        |  +-----------------------+                         
        |  |                       +-- Hyridella_australis   
        |  |                                                 
        +--+  +----------------------- Echyridella           
           |  |                                              
           |  |              +-------- Velesunio__1 **          
           +--+  +-----------+                               
              |  |           |  +----- Alathyria_jacksoni__1 ** 
              |  |           +--+                            
              |  |              |  +-- Velesunio_ambiguus **    
              +--+              +--+                         
                 |                 +-- Alathyria_profuga **     
                 |                                           
                 |  +----------------- Velesunio__3 **          
                 |  |                                        
                 +--+  +-------------- Microdontia **           
                    |  |                                     
                    +--+           +-- Lortiella **             
                       |  +--------+                         
                       |  |        +-- Velesunio_angasi **      
                       +--+                                  
                          |  +-------- Alathyria_pertexta__2 ** 
                          +--+                               
                             |  +----- Velesunio__2 **          
                             +--+                            
                                |  +-- Alathyria_jacksoni__2 ** 
                                +--+                         
                                   +-- Alathyria_pertexta__1 ** 

Graf et al., 2015 [Publ 3619], fig. 1

iTtTbiriSoTroTroSiD
7755137136
  +-------------- Hyriinae          
  |                                 
--+  +----------- Westralunio **       
  |  |                              
  |  |     +----- Alathyria_jacksoni **
  +--+  +--+                        
     |  |  |  +-- Velesunio **         
     |  |  +--+                     
     +--+     +-- Alathyria_profuga ** 
        |                           
        |  +----- Alathyria_pertexta **
        +--+                        
           |  +-- Microdontia **       
           +--+                     
              +-- Lortiella **         

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

iTtTbiriSoTroTroSiD
7141251937139
           +-- Westralunio **       
  +--------+                     
  |        +-- Hyriinae          
  |                              
--+     +----- Alathyria_jacksoni **
  |  +--+                        
  |  |  |  +-- Alathyria_profuga ** 
  |  |  +--+                     
  +--+     +-- Velesunio **         
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **       

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

iTtTbiriSoTroTroSiD
7141251917141
           +-- Westralunio **       
  +--------+                     
  |        +-- Hyriinae          
  |                              
--+     +----- Velesunio **         
  |  +--+                        
  |  |  |  +-- Alathyria_jacksoni **
  |  |  +--+                     
  +--+     +-- Alathyria_profuga ** 
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **       

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

iTtTbiriSoTroTroSiD
714951937138
  +----------- Echyridella       
  |                              
  +----------- Westralunio **       
  |                              
  +----------- Hyridellini       
  |                              
  |     +----- Rhipidodontini    
  +-----+                        
--+     |  +-- Castaliini        
  |     +--+                     
  |        +-- Hyriini           
  |                              
  |     +----- Velesunio **         
  |  +--+                        
  |  |  |  +-- Alathyria_jacksoni **
  |  |  +--+                     
  +--+     +-- Alathyria_profuga ** 
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **       

Santos-Neto et al., 2016 [Publ 3739], fig. 4

iTtTbiriSoTroTroSiD
6131151413131.5
                    +----- Diplodon_suavidicus__2
  +-----------------+                            
  |                 |  +-- Hyriini               
  |                 +--+                         
  |                    +-- Castaliini            
  |                                              
  |              +-------- Westralunio **           
--+              |                               
  |  +-----------+     +-- Hyridella_drapeta     
  |  |           |  +--+                         
  |  |           |  |  +-- Hyridella_depressa    
  |  |           +--+                            
  |  |              |  +-- Diplodon_suavidicus__1
  +--+              +--+                         
     |                 +-- Diplodon_gratus       
     |                                           
     |  +----------------- Echyridella           
     |  |                                        
     +--+              +-- Cucumerunio           
        |  +-----------+                         
        |  |           +-- Hyridella_australis   
        +--+                                     
           |  +----------- Velesunio_ambiguus **    
           +--+                                  
              |  +-------- Alathyria **             
              +--+                               
                 |  +----- Velesunio_angasi **      
                 +--+                            
                    |  +-- Lortiella **             
                    +--+                         
                       +-- Microdontia **           

Bolotov et al., 2016 [Publ 3743], fig. 4

iTtTbiriSoTroTroSiD
6644243139
  +----------- Hyriinae          
  |                              
  |     +----- Microdontia **       
--+  +--+                        
  |  |  |  +-- Alathyria_pertexta **
  |  |  +--+                     
  +--+     +-- Lortiella **         
     |                           
     |  +----- Velesunio **         
     +--+                        
        |  +-- Alathyria_jacksoni **
        +--+                     
           +-- Alathyria_profuga ** 

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

iTtTbiriSoTroTroSiD
66441323139
  +----------- Hyriinae          
  |                              
  |     +----- Velesunio **         
--+  +--+                        
  |  |  |  +-- Alathyria_jacksoni **
  |  |  +--+                     
  +--+     +-- Alathyria_profuga ** 
     |                           
     |  +----- Microdontia **       
     +--+                        
        |  +-- Alathyria_pertexta **
        +--+                     
           +-- Lortiella **         

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

iTtTbiriSoTroTroSiD
66441323139
  +----------- Hyriinae          
  |                              
  |     +----- Alathyria_pertexta **
--+  +--+                        
  |  |  |  +-- Microdontia **       
  |  |  +--+                     
  +--+     +-- Lortiella **         
     |                           
     |  +----- Velesunio **         
     +--+                        
        |  +-- Alathyria_jacksoni **
        +--+                     
           +-- Alathyria_profuga ** 

Santos-Neto et al., 2016 [Publ 3739], fig. 2a_b

iTtTbiriSoTroTroSiD
6192051413122.5
                 +----- Diplodon_suavidicus__1
  +--------------+                            
  |              |  +-- Hyriini               
  |              +--+                         
  |                 +-- Castaliini            
  |                                           
  |           +-------- Westralunio **           
  |           |                               
  +-----------+     +-- Hyridella_drapeta     
  |           |  +--+                         
--+           |  |  +-- Hyridella_depressa    
  |           +--+                            
  |              |  +-- Diplodon_gratus       
  |              +--+                         
  |                 +-- Diplodon_suavidicus__2
  |                                           
  |                 +-- Hyridella_australis   
  |  +--------------+                         
  |  |              +-- Cucumerunio           
  +--+                                        
     |  +-------------- Echyridella           
     +--+                                     
        |  +----------- Velesunio_ambiguus **    
        +--+                                  
           |  +-------- Microdontia **           
           +--+                               
              |  +----- Lortiella **             
              +--+                            
                 |  +-- Alathyria **             
                 +--+                         
                    +-- Velesunio_angasi **      

Bolotov et al., 2016 [Publ 3743], fig. 2

iTtTbiriSoTroTroSiD
6644243139
  +----------- Hyriinae          
  |                              
  |     +----- Microdontia **       
--+  +--+                        
  |  |  |  +-- Lortiella **         
  |  |  +--+                     
  +--+     +-- Alathyria_pertexta **
     |                           
     |  +----- Velesunio **         
     +--+                        
        |  +-- Alathyria_jacksoni **
        +--+                     
           +-- Alathyria_profuga ** 

Walker et al., 2006 [Publ 2666], fig. 6

iTtTbiriSoTroTroSiD
5543315130
  +----------------- Castaliini         
  |                                     
  +----------------- Rhipidodontini     
--+                                     
  |  +-------------- Hyridella_depressa 
  |  |                                  
  |  +-------------- Hyridella_australis
  +--+                                  
     |  +----------- Echyridella        
     |  |                               
     |  |        +-- Velesunio_ambiguus ** 
     +--+  +-----+                      
        |  |     +-- Velesunio__3 **       
        |  |                            
        +--+  +----- Velesunio__2 **       
           |  |                         
           |  |  +-- Alathyria **          
           +--+--+                      
              |  +-- Velesunio__1 **       
              |                         
              |  +-- Velesunio_angasi **   
              +--+                      
                 +-- Lortiella **          

Marshall et al., 2014 [Publ 3593], fig. 1

iTtTbiriSoTroTroSiD
3312117134.5
  +----------- Echyridella       
  |                              
--+        +-- Hyridellini       
  |  +-----+                     
  |  |     +-- Rhipidodontini    
  +--+                           
     |  +----- Velesunio_ambiguus **
     +--+                        
        |  +-- Velesunio **         
        +--+                     
           +-- Lortiella **         

Baker et al., 2004 [Publ 2685], fig. 2-3

iTtTbiriSoTroTroSiD
334265132
              +----- Hyridella_depressa 
  +-----------+                         
  |           |  +-- Hyridella_australis
  |           +--+                      
  |              +-- Rhipidodontini     
--+                                     
  |              +-- Echyridella        
  |  +-----------+                      
  |  |           +-- Castaliini         
  |  |                                  
  +--+           +-- Velesunio__3 **       
     |  +--------+                      
     |  |        +-- Velesunio__4 **       
     +--+                               
        |  +-------- Velesunio__2 **       
        +--+                            
           |  +----- Velesunio_angasi **   
           +--+                         
              |  +-- Velesunio__1 **       
              +--+                      
                 +-- Alathyria **          

Klunzinger et al., 2021 [Publ 4086], fig. 2

iTtTbiriSoTroTroSiD
342231132
  +----- Westralunio **
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Hyriinae   

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

iTtTbiriSoTroTroSiD
3312436133
  +-------- Hyriinae          
--+                           
  |  +----- Velesunio_ambiguus **
  +--+                        
     |  +-- Velesunio_angasi **  
     +--+                     
        +-- Lortiella **         

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

iTtTbiriSoTroTroSiD
3312446133
  +-------- Hyriinae          
--+                           
  |  +----- Velesunio_ambiguus **
  +--+                        
     |  +-- Velesunio_angasi **  
     +--+                     
        +-- Lortiella **         

Klunzinger et al., 2022 [Publ 4259], fig. 1

iTtTbiriSoTroTroSiD
331211133
  +----- Hyriinae   
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Westralunio **

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

iTtTbiriSoTroTroSiD
3312446133
  +-------- Hyriinae          
--+                           
  |  +----- Velesunio_ambiguus **
  +--+                        
     |  +-- Velesunio_angasi **  
     +--+                     
        +-- Lortiella **         

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

iTtTbiriSoTroTroSiD
3312436133
  +-------- Hyriinae          
--+                           
  |  +----- Velesunio_ambiguus **
  +--+                        
     |  +-- Velesunio_angasi **  
     +--+                     
        +-- Lortiella **         

Hoeh et al., 2001 [Publ 2652], fig. 14f2

iTtTbiriSoTroTroSiD
2502293135
  +----- Rhipidodontini
  |                    
  +----- Echyridella   
--+                    
  +----- Lortiella **     
  |                    
  |  +-- Castaliini    
  +--+                 
     +-- Velesunio **     

Baker et al., 2003 [Publ 2690], fig. 5

iTtTbiriSoTroTroSiD
2201651--
           +-- Hyridella_australis
  +--------+                      
  |        +-- Rhipidodontini     
--+                               
  |  +-------- Hyridella_depressa 
  +--+                            
     |  +----- Velesunio **          
     +--+                         
        |  +-- Echyridella        
        +--+                      
           +-- Castaliini         

Hoeh et al., 2002 [Publ 2653], fig. 2

iTtTbiriSoTroTroSiD
22022931--
        +-- Castaliini    
  +-----+                 
  |     +-- Rhipidodontini
--+                       
  |  +----- Echyridella   
  +--+                    
     |  +-- Velesunio **     
     +--+                 
        +-- Lortiella **     

Roe & Hoeh, 2003 [Publ 2655], fig. 4f2

iTtTbiriSoTroTroSiD
2502293136
     +-- Castaliini    
  +--+                 
  |  +-- Velesunio **     
--+                    
  |  +-- Echyridella   
  |  |                 
  +--+-- Lortiella **     
     |                 
     +-- Rhipidodontini

Bogan & Hoeh, 2000 [Publ 2650], fig. 1

iTtTbiriSoTroTroSiD
22023231--
        +-- Castaliini    
  +-----+                 
  |     +-- Rhipidodontini
--+                       
  |  +----- Echyridella   
  +--+                    
     |  +-- Velesunio **     
     +--+                 
        +-- Lortiella **     

Hoeh et al., 2001 [Publ 2652], fig. 14f4

iTtTbiriSoTroTroSiD
22022931--
        +-- Castaliini    
  +-----+                 
  |     +-- Rhipidodontini
--+                       
  |  +----- Echyridella   
  +--+                    
     |  +-- Lortiella **     
     +--+                 
        +-- Velesunio **     

Roe & Hoeh, 2003 [Publ 2655], fig. 4f3

iTtTbiriSoTroTroSiD
22022931--
        +-- Castaliini    
  +-----+                 
  |     +-- Rhipidodontini
--+                       
  |  +----- Echyridella   
  +--+                    
     |  +-- Velesunio **     
     +--+                 
        +-- Lortiella **     

Guerra et al., 2019 [Publ 4168], fig. 2

iTtTbiriSoTroTroSiD
11014041--
  +-- Westralunio **
--+              
  +-- Hyriinae   

Plazzi et al., 2011 [Publ 3676], fig. 5

iTtTbiriSoTroTroSiD
11011700--
-- Alathyria_jacksoni **

Sonowal et al., 2021 [Publ 4221], fig. 4

iTtTbiriSoTroTroSiD
11011521--
  +----- Hyriini    
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Hyridellini

Sonowal et al., 2021 [Publ 4221], fig. 3a

iTtTbiriSoTroTroSiD
11011421--
  +-- Hyriini    
  |              
--+-- Hyridellini
  |              
  +-- Velesunio **  

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
11011731--
  +-- Velesunio **
--+            
  +-- Hyriinae 

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

iTtTbiriSoTroTroSiD
110112111--
  +-- Hyriinae 
--+            
  +-- Velesunio **

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

iTtTbiriSoTroTroSiD
110114041--
  +-- Westralunio **
--+              
  +-- Hyriinae   

González & Giribet, 2015 [Publ 3632], fig. 6

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

Lopes-Lima et al., 2018 [Publ 3829], fig. 3

iTtTbiriSoTroTroSiD
11012721--
  +----- Hyriini    
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Hyridellini

Combosch et al., 2017 [Publ 3806], fig. 1

iTtTbiriSoTroTroSiD
11011421--
  +----- Hyriini    
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Hyridellini

Huang et al., 2018 [Publ 3828], fig. 3_5

iTtTbiriSoTroTroSiD
11012821--
  +----- Hyriini    
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Hyridellini

Wu et al., 2020 [Publ 4121], fig. 2

iTtTbiriSoTroTroSiD
11013321--
  +----- Hyriini    
--+                 
  |  +-- Hyridellini
  +--+              
     +-- Velesunio **  

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

iTtTbiriSoTroTroSiD
11013000--
-- Velesunio_ambiguus **

Combosch et al., 2017 [Publ 3806], fig. 2

iTtTbiriSoTroTroSiD
11011421--
  +-- Velesunio **
--+            
  +-- Hyriinae 

González & Giribet, 2015 [Publ 3632], fig. 2

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
11014611--
  +-- Hyriinae   
--+              
  +-- Westralunio **

Sano et al., 2020 [Publ 4138], fig. 2

iTtTbiriSoTroTroSiD
11016721--
  +----- Hyriini    
--+                 
  |  +-- Hyridellini
  +--+              
     +-- Velesunio **  

González & Giribet, 2015 [Publ 3632], fig. 3

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

Elderkin et al., 2016 [Publ 3693], fig. 2

iTtTbiriSoTroTroSiD
1101200--
-- Velesunio_ambiguus **

Bieler et al., 2014 [Publ 3622], fig. 26_27_29_30

iTtTbiriSoTroTroSiD
1101400--
-- Velesunio_ambiguus **

González & Giribet, 2015 [Publ 3632], fig. 4

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
11017911--
  +-- Hyriinae 
--+            
  +-- Velesunio **

Sonowal et al., 2021 [Publ 4221], fig. 3b

iTtTbiriSoTroTroSiD
11011421--
  +-- Hyriini    
  |              
--+-- Velesunio **  
  |              
  +-- Hyridellini

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

iTtTbiriSoTroTroSiD
11019021--
  +-- Alathyria **
--+            
  +-- Hyriinae 

Wu et al., 2020 [Publ 4121], fig. S1

iTtTbiriSoTroTroSiD
11013321--
  +----- Hyriini    
--+                 
  |  +-- Hyridellini
  +--+              
     +-- Velesunio **  

Sano et al., 2020 [Publ 4138], fig. 3

iTtTbiriSoTroTroSiD
11016721--
  +----- Hyriini    
--+                 
  |  +-- Hyridellini
  +--+              
     +-- Velesunio **  

Guerra et al., 2019 [Publ 4168], fig. 1-M

iTtTbiriSoTroTroSiD
11012111--
  +-- Hyriinae   
--+              
  +-- Westralunio **

Sano et al., 2020 [Publ 4138], fig. 4

iTtTbiriSoTroTroSiD
11016721--
  +----- Hyriini    
--+                 
  |  +-- Hyridellini
  +--+              
     +-- Velesunio **  

Guerra et al., 2019 [Publ 4168], fig. 2-M

iTtTbiriSoTroTroSiD
11012111--
  +-- Hyriinae   
--+              
  +-- Westralunio **

Guerra et al., 2019 [Publ 4168], fig. 1

iTtTbiriSoTroTroSiD
11014041--
        +-- Echyridella   
  +-----+                 
  |     +-- Westralunio **   
--+                       
  |  +----- Rhipidodontini
  +--+                    
     |  +-- Castaliini    
     +--+                 
        +-- Hyriini       

Lopes-Lima et al., 2018 [Publ 3829], fig. 1

iTtTbiriSoTroTroSiD
11012721--
  +----- Hyriini    
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Hyridellini

Bieler et al., 2014 [Publ 3622], fig. 28

iTtTbiriSoTroTroSiD
1101400--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
11013531--
  +-- Velesunio **
--+            
  +-- Hyriinae 

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

iTtTbiriSoTroTroSiD
11013531--
  +-- Velesunio **
--+            
  +-- Hyriinae 

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

iTtTbiriSoTroTroSiD
11013931--
  +-- Velesunio **
--+            
  +-- Hyriinae 

Graf & Ó Foighil, 2000 [Publ 2649], fig. 1_2

iTtTbiriSoTroTroSiD
11011051--
  +-- Velesunio **
--+            
  +-- Hyriinae 

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

iTtTbiriSoTroTroSiD
11019021--
  +-- Alathyria **
--+            
  +-- Hyriinae 

Graf, 2000 [Publ 2647], fig. 1

iTtTbiriSoTroTroSiD
11011631--
  +-------- Castaliini    
--+                       
  |  +----- Rhipidodontini
  +--+                    
     |  +-- Hyridellini   
     +--+                 
        +-- Velesunio **     
 
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