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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 | 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 62 cladograms in 37 publications
maximum number of terminals in an analysis = 7 (mean = 2.31, median = 1)
maximum number of subtaxa (genus) = 5 (mean = 1.84, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 26
maximum number of included terminals = 7, (mean = 4.12 median = 3)
maximum number of subtaxa (genus) = 5 (mean = 3, median = 2)
proportion of cladograms that support basic monophyly = 0.69

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 25
(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 = 7 (mean = 4.2, median = 3)
maximum number of subtaxa (genus) = 5 (mean = 3.08, median = 2)
proportion of cladograms that support rigorous monophyly = 0.68

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

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

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

iTtTbiriSoTroTroSiD
619205141310
                 +----- 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 **

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

iTtTbiriSoTroTroSiD
7141251947126.5
           +-- Westralunio **       
  +--------+                     
  |        +-- Hyriinae          
  |                              
--+     +----- Alathyria_jacksoni **
  |  +--+                        
  |  |  |  +-- Alathyria_profuga ** 
  |  |  +--+                     
  +--+     +-- Velesunio **         
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **

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

iTtTbiriSoTroTroSiD
7755137124.5
  +-------------- Hyriinae          
  |                                 
--+  +----------- Westralunio **       
  |  |                              
  |  |     +----- Alathyria_jacksoni **
  +--+  +--+                        
     |  |  |  +-- Velesunio **         
     |  |  +--+                     
     +--+     +-- Alathyria_profuga ** 
        |                           
        |  +----- Alathyria_pertexta **
        +--+                        
           |  +-- Microdontia **       
           +--+                     
              +-- Lortiella **

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

iTtTbiriSoTroTroSiD
714951947123.5
  +----------- Echyridella       
  |                              
  +----------- Westralunio **       
  |                              
  +----------- Hyridellini       
  |                              
  |     +----- Rhipidodontini    
  +-----+                        
--+     |  +-- Castaliini        
  |     +--+                     
  |        +-- Hyriini           
  |                              
  |     +----- Velesunio **         
  |  +--+                        
  |  |  |  +-- Alathyria_jacksoni **
  |  |  +--+                     
  +--+     +-- Alathyria_profuga ** 
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **

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

iTtTbiriSoTroTroSiD
7141251927126.5
           +-- Westralunio **       
  +--------+                     
  |        +-- Hyriinae          
  |                              
--+     +----- Velesunio **         
  |  +--+                        
  |  |  |  +-- Alathyria_jacksoni **
  |  |  +--+                     
  +--+     +-- Alathyria_profuga ** 
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **

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

iTtTbiriSoTroTroSiD
6644243125.5
  +----------- Hyriinae          
  |                              
  |     +----- Microdontia **       
--+  +--+                        
  |  |  |  +-- Alathyria_pertexta **
  |  |  +--+                     
  +--+     +-- Lortiella **         
     |                           
     |  +----- Velesunio **         
     +--+                        
        |  +-- Alathyria_jacksoni **
        +--+                     
           +-- Alathyria_profuga **

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

iTtTbiriSoTroTroSiD
66441333125.5
  +----------- Hyriinae          
  |                              
  |     +----- Velesunio **         
--+  +--+                        
  |  |  |  +-- Alathyria_jacksoni **
  |  |  +--+                     
  +--+     +-- Alathyria_profuga ** 
     |                           
     |  +----- Microdontia **       
     +--+                        
        |  +-- Alathyria_pertexta **
        +--+                     
           +-- Lortiella **

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

iTtTbiriSoTroTroSiD
6644243125.5
  +----------- Hyriinae          
  |                              
  |     +----- Microdontia **       
--+  +--+                        
  |  |  |  +-- Lortiella **         
  |  |  +--+                     
  +--+     +-- Alathyria_pertexta **
     |                           
     |  +----- Velesunio **         
     +--+                        
        |  +-- Alathyria_jacksoni **
        +--+                     
           +-- Alathyria_profuga **

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

iTtTbiriSoTroTroSiD
66441333125.5
  +----------- Hyriinae          
  |                              
  |     +----- Alathyria_pertexta **
--+  +--+                        
  |  |  |  +-- Microdontia **       
  |  |  +--+                     
  +--+     +-- Lortiella **         
     |                           
     |  +----- Velesunio **         
     +--+                        
        |  +-- Alathyria_jacksoni **
        +--+                     
           +-- Alathyria_profuga **

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

iTtTbiriSoTroTroSiD
6131151413116
                    +----- 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 **

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

iTtTbiriSoTroTroSiD
5543315122.5
  +----------------- Castaliini         
  |                                     
  +----------------- Rhipidodontini     
--+                                     
  |  +-------------- Hyridella_depressa 
  |  |                                  
  |  +-------------- Hyridella_australis
  +--+                                  
     |  +----------- Echyridella        
     |  |                               
     |  |        +-- Velesunio_ambiguus ** 
     +--+  +-----+                      
        |  |     +-- Velesunio__3 **       
        |  |                            
        +--+  +----- Velesunio__2 **       
           |  |                         
           |  |  +-- Alathyria **          
           +--+--+                      
              |  +-- Velesunio__1 **       
              |                         
              |  +-- Velesunio_angasi **   
              +--+                      
                 +-- Lortiella **

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

iTtTbiriSoTroTroSiD
3312446120.5
  +-------- Hyriinae          
--+                           
  |  +----- Velesunio_ambiguus **
  +--+                        
     |  +-- Velesunio_angasi **  
     +--+                     
        +-- Lortiella **

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

iTtTbiriSoTroTroSiD
3312117122
  +----------- Echyridella       
  |                              
--+        +-- Hyridellini       
  |  +-----+                     
  |  |     +-- Rhipidodontini    
  +--+                           
     |  +----- Velesunio_ambiguus **
     +--+                        
        |  +-- Velesunio **         
        +--+                     
           +-- Lortiella **

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

iTtTbiriSoTroTroSiD
3312436120.5
  +-------- Hyriinae          
--+                           
  |  +----- Velesunio_ambiguus **
  +--+                        
     |  +-- Velesunio_angasi **  
     +--+                     
        +-- Lortiella **

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

iTtTbiriSoTroTroSiD
331211122
  +----- Hyriinae   
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Westralunio **

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

iTtTbiriSoTroTroSiD
342231121
  +----- Westralunio **
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Hyriinae

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

iTtTbiriSoTroTroSiD
3312446120.5
  +-------- Hyriinae          
--+                           
  |  +----- Velesunio_ambiguus **
  +--+                        
     |  +-- Velesunio_angasi **  
     +--+                     
        +-- Lortiella **

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

iTtTbiriSoTroTroSiD
3312436120.5
  +-------- Hyriinae          
--+                           
  |  +----- Velesunio_ambiguus **
  +--+                        
     |  +-- Velesunio_angasi **  
     +--+                     
        +-- Lortiella **

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

iTtTbiriSoTroTroSiD
334265122
              +----- Hyridella_depressa 
  +-----------+                         
  |           |  +-- Hyridella_australis
  |           +--+                      
  |              +-- Rhipidodontini     
--+                                     
  |              +-- Echyridella        
  |  +-----------+                      
  |  |           +-- Castaliini         
  |  |                                  
  +--+           +-- Velesunio__3 **       
     |  +--------+                      
     |  |        +-- Velesunio__4 **       
     +--+                               
        |  +-------- Velesunio__2 **       
        +--+                            
           |  +----- Velesunio_angasi **   
           +--+                         
              |  +-- Velesunio__1 **       
              +--+                      
                 +-- Alathyria **

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

iTtTbiriSoTroTroSiD
2502293122.5
     +-- Castaliini    
  +--+                 
  |  +-- Velesunio **     
--+                    
  |  +-- Echyridella   
  |  |                 
  +--+-- Lortiella **     
     |                 
     +-- Rhipidodontini

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
2502293121.5
  +----- Rhipidodontini
  |                    
  +----- Echyridella   
--+                    
  +----- Lortiella **     
  |                    
  |  +-- Castaliini    
  +--+                 
     +-- Velesunio **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1101400--
-- Velesunio_ambiguus **

Graf, 2000 [Publ 2647], fig. 1

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
11011700--
-- Alathyria_jacksoni **

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

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

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

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

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

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

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

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

iTtTbiriSoTroTroSiD
1101200--
-- Velesunio_ambiguus **

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

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

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

iTtTbiriSoTroTroSiD
1101400--
-- Velesunio_ambiguus **

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

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

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

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

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

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

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

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

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

Graf, 2002 [Publ 2651], fig. 2

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
11013000--
-- Velesunio_ambiguus **
 
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