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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

family HYRIIDAE Swainson, 1840: cladomics
     2 genera, 2 subfamilies

Cladomics Summary

number of species = 93 (132 total)
number of subtaxa (rank: subfamily) = 4
analyzed in 102 cladograms in 54 publications
maximum number of terminals in an analysis = 19 (mean = 3.84, median = 3)
maximum number of subtaxa (subfamily) = 2 (mean = 1.52, median = 2)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 58
maximum number of included terminals = 19, (mean = 6 median = 4.5)
maximum number of subtaxa (subfamily) = 2 (mean = 1.91, median = 2)
proportion of cladograms that support basic monophyly = 1

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

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

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

iTtTbiriSoTroTroSiD
88921000
     +----- Hyridella_depressa ** 
  +--+                         
  |  |  +-- Hyridella_australis **
  |  +--+                      
--+     +-- Rhipidodontini **     
  |                            
  |  +----- Velesunioninae **     
  +--+                         
     |  +-- Echyridella **        
     +--+                      
        +-- Castaliini **

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

iTtTbiriSoTroTroSiD
191921210025
                    +----- 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 **

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

iTtTbiriSoTroTroSiD
191920210024
                 +----- 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. S3

iTtTbiriSoTroTroSiD
14141221850021.5
           +-- Westralunio **       
  +--------+                     
  |        +-- Hyriinae **          
  |                              
--+     +----- Velesunio **         
  |  +--+                        
  |  |  |  +-- Alathyria_jacksoni **
  |  |  +--+                     
  +--+     +-- Alathyria_profuga ** 
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **

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

iTtTbiriSoTroTroSiD
14141221870021.5
           +-- Westralunio **       
  +--------+                     
  |        +-- Hyriinae **          
  |                              
--+     +----- Alathyria_jacksoni **
  |  +--+                        
  |  |  |  +-- Alathyria_profuga ** 
  |  |  +--+                     
  +--+     +-- Velesunio **         
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **

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

iTtTbiriSoTroTroSiD
141412260021.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
1414921870019.5
  +----------- Echyridella **       
  |                              
  +----------- Westralunio **       
  |                              
  +----------- Hyridellini **       
  |                              
  |     +----- Rhipidodontini **    
  +-----+                        
--+     |  +-- Castaliini **        
  |     +--+                     
  |        +-- Hyriini **           
  |                              
  |     +----- Velesunio **         
  |  +--+                        
  |  |  |  +-- Alathyria_jacksoni **
  |  |  +--+                     
  +--+     +-- Alathyria_profuga ** 
     |                           
     |  +----- Alathyria_pertexta **
     +--+                        
        |  +-- Lortiella **         
        +--+                     
           +-- Microdontia **

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

iTtTbiriSoTroTroSiD
10108240015
  +-------- Echyridella **   
--+                       
  |  +----- Velesunioninae **
  +--+                    
     |  +-- Hyridellini **   
     +--+                 
        +-- Rhipidodontini **

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

iTtTbiriSoTroTroSiD
10107226009
  +-------- Castaliini **         
  |                            
  +-------- Rhipidodontini **     
--+                            
  |  +----- Hyridella_depressa ** 
  |  |                         
  +--+----- Hyridella_australis **
     |                         
     |  +-- Echyridella **        
     +--+                      
        +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
9972380015.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
9972370015.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
9972210016.5
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
9972370015.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
99721300016.5
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
9972210016.5
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
9972380015.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
99721300016.5
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
77521008
           +-- Hyridella_australis **
  +--------+                      
  |        +-- Rhipidodontini **     
--+                               
  |  +-------- Hyridella_depressa ** 
  +--+                            
     |  +----- Velesunioninae **     
     +--+                         
        |  +-- Echyridella **        
        +--+                      
           +-- Castaliini **

Santos-Neto et al., 2016 [Publ 3739], fig. 3_5

iTtTbiriSoTroTroSiD
776100017
1

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

iTtTbiriSoTroTroSiD
665250016
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
5532360015
        +-- Echyridella **   
  +-----+                 
  |     +-- Velesunioninae **
--+                       
  |  +----- Rhipidodontini **
  +--+                    
     |  +-- Castaliini **    
     +--+                 
        +-- Hyriini **

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

iTtTbiriSoTroTroSiD
5522260012
     +-- Castaliini **    
  +--+                 
  |  +-- Velesunio **     
--+                    
  |  +-- Echyridella **   
  |  |                 
  +--+-- Lortiella **     
     |                 
     +-- Rhipidodontini **

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

iTtTbiriSoTroTroSiD
5532290012
     +-- Castaliini **    
  +--+                 
  |  +-- Rhipidodontini **
--+                    
  |  +-- Echyridella **   
  +--+                 
     +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
5532260012
     +-- Castaliini **    
  +--+                 
  |  +-- Rhipidodontini **
--+                    
  |  +-- Echyridella **   
  +--+                 
     +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
5532260012
     +-- Castaliini **    
  +--+                 
  |  +-- Rhipidodontini **
--+                    
  |  +-- Echyridella **   
  +--+                 
     +-- Velesunioninae **

Santos-Neto et al., 2018 [Publ 3888], fig. 5

iTtTbiriSoTroTroSiD
553100014
1

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

iTtTbiriSoTroTroSiD
5532260012
     +-- Castaliini **    
  +--+                 
  |  +-- Rhipidodontini **
--+                    
  |  +-- Echyridella **   
  +--+                 
     +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
5532360015
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
5512260011
  +----- Rhipidodontini **
  |                    
  +----- Echyridella **   
--+                    
  +----- Lortiella **     
  |                    
  |  +-- Castaliini **    
  +--+                 
     +-- Velesunio **

Graf, 2000 [Publ 2647], fig. 1

iTtTbiriSoTroTroSiD
4422130012.5
  +-------- Castaliini **    
--+                       
  |  +----- Rhipidodontini **
  +--+                    
     |  +-- Hyridellini **   
     +--+                 
        +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
4422320012.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
4422140013
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
4422320012.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
4422360012.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
442200012.5
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
442220012.5
  +----- Westralunio **
--+                 
  |  +-- Velesunio **  
  +--+              
     +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
3312310010.5
  +----- Hyriini **       
--+                    
  |  +-- Hyridellini **   
  +--+                 
     +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
3312250010.5
  +----- Hyriini **       
--+                    
  |  +-- Velesunioninae **
  +--+                 
     +-- Hyridellini **

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

iTtTbiriSoTroTroSiD
3312880012
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
3312650010.5
  +----- Hyriini **       
--+                    
  |  +-- Hyridellini **   
  +--+                 
     +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
3312120010.5
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
3312130010.5
  +----- Hyriini **       
--+                    
  |  +-- Velesunioninae **
  +--+                 
     +-- Hyridellini **

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

iTtTbiriSoTroTroSiD
3312880012
  +-- Velesunioninae **
--+                 
  +-- Hyriinae **

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

iTtTbiriSoTroTroSiD
3302120010.5
  +-- Hyriini **       
  |                 
--+-- Hyridellini **   
  |                 
  +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
3312650010.5
  +----- Hyriini **       
--+                    
  |  +-- Hyridellini **   
  +--+                 
     +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
3311280012
1

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

iTtTbiriSoTroTroSiD
3312310010.5
  +----- Hyriini **       
--+                    
  |  +-- Hyridellini **   
  +--+                 
     +-- Velesunioninae **

Olivera-Hyde et al., 2020 [Publ 4249], fig. 10

iTtTbiriSoTroTroSiD
331100012
1

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

iTtTbiriSoTroTroSiD
3312250010.5
  +----- Hyriini **       
--+                    
  |  +-- Velesunioninae **
  +--+                 
     +-- Hyridellini **

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

iTtTbiriSoTroTroSiD
3312650010.5
  +----- Hyriini **       
--+                    
  |  +-- Hyridellini **   
  +--+                 
     +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
3312120010.5
  +----- Hyriini **       
--+                    
  |  +-- Velesunioninae **
  +--+                 
     +-- Hyridellini **

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

iTtTbiriSoTroTroSiD
3312260010.5
  +----- Hyriini **       
--+                    
  |  +-- Velesunioninae **
  +--+                 
     +-- Hyridellini **

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

iTtTbiriSoTroTroSiD
3302120010.5
  +-- Hyriini **       
  |                 
--+-- Velesunioninae **
  |                 
  +-- Hyridellini **

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

iTtTbiriSoTroTroSiD
22022000--
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
22027800--
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

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

iTtTbiriSoTroTroSiD
22022000--
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

Chen et al., 2025 [Publ 4455], fig. 1

iTtTbiriSoTroTroSiD
22012400--
1

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

iTtTbiriSoTroTroSiD
220212000--
  +-- Hyriinae **      
--+                 
  +-- Velesunioninae **

Guerra et al., 2017 [Publ 3772], fig. 2a

iTtTbiriSoTroTroSiD
11012400--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
1101400--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
1101200--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
11019500--
-- Echyridella_menziesii **

Huang et al., 2019 [Publ 3843], fig. 3-M

iTtTbiriSoTroTroSiD
11012200--
-- Echyridella_menziesii **

Huang et al., 2018 [Publ 3828], fig. 6-M

iTtTbiriSoTroTroSiD
11011600--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
11018700--
-- Echyridella_menziesii **

Walker et al., 2006 [Publ 2666], fig. 3

iTtTbiriSoTroTroSiD
11011400--
-- Hyridella **

Chase et al., 2018 [Publ 3861], fig. 1-M

iTtTbiriSoTroTroSiD
11011000--
-- Echyridella_menziesii **

Guerra et al., 2017 [Publ 3772], fig. 2b-M

iTtTbiriSoTroTroSiD
1101700--
-- Echyridella_menziesii **

Guerra et al., 2017 [Publ 3772], fig. 2b

iTtTbiriSoTroTroSiD
11012400--
-- Echyridella_menziesii **

Chase et al., 2018 [Publ 3847], fig. 4

iTtTbiriSoTroTroSiD
11013700--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
11019100--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
11018500--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
1101400--
-- Velesunio_ambiguus **

Walker et al., 2006 [Publ 2666], fig. 2

iTtTbiriSoTroTroSiD
11011400--
-- Hyridella **

Wu et al., 2019 [Publ 3864], fig. 3

iTtTbiriSoTroTroSiD
11014500--
-- Echyridella_menziesii **

Wu et al., 2021 [Publ 4366], fig. 4

iTtTbiriSoTroTroSiD
11016800--
-- Echyridella_menziesii **

Graf & Ó Foighil, 2000 [Publ 2648], fig. 2B

iTtTbiriSoTroTroSiD
11012900--
-- Hyridella_depressa **

Wu et al., 2022 [Publ 4238], fig. 3

iTtTbiriSoTroTroSiD
11017300--
-- Echyridella_menziesii **

Huang et al., 2018 [Publ 3828], fig. 6

iTtTbiriSoTroTroSiD
11014100--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
11019100--
-- Echyridella_menziesii **

Wu et al., 2020 [Publ 4121], fig. 3

iTtTbiriSoTroTroSiD
11014800--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

Huang et al., 2019 [Publ 3843], fig. 3

iTtTbiriSoTroTroSiD
11015500--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
11015600--
-- Hyridella_australis **

Graf & Ó Foighil, 2000 [Publ 2648], fig. 2A

iTtTbiriSoTroTroSiD
11012900--
-- Hyridella_depressa **

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

iTtTbiriSoTroTroSiD
1101500--
-- Velesunio_ambiguus **

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

iTtTbiriSoTroTroSiD
11011700--
-- Alathyria_jacksoni **

Hoeh et al., 2002 [Publ 2673], fig. 1-F

iTtTbiriSoTroTroSiD
1101800--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
11011300--
-- Echyridella_menziesii **

Wu et al., 2021 [Publ 4366], fig. 3-ML

iTtTbiriSoTroTroSiD
11017300--
-- Echyridella_menziesii **

Chase et al., 2018 [Publ 3861], fig. 1

iTtTbiriSoTroTroSiD
11011000--
-- Echyridella_menziesii **

Hoeh et al., 2002 [Publ 2673], fig. 1-M

iTtTbiriSoTroTroSiD
1101800--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
11019500--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
11015600--
-- Hyridella_australis **

Wu et al., 2021 [Publ 4366], fig. 3-BI

iTtTbiriSoTroTroSiD
11017300--
-- Echyridella_menziesii **

Guerra et al., 2017 [Publ 3772], fig. 2a-M

iTtTbiriSoTroTroSiD
1101700--
-- Echyridella_menziesii **

Chase et al., 2018 [Publ 3847], fig. 5

iTtTbiriSoTroTroSiD
1101900--
-- Echyridella_menziesii **

Walker et al., 2006 [Publ 2666], fig. 4

iTtTbiriSoTroTroSiD
11011400--
-- Hyridella **

Breton et al., 2011 [Publ 3376], fig. 3

iTtTbiriSoTroTroSiD
11013600--
-- Echyridella_menziesii **

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

iTtTbiriSoTroTroSiD
11013000--
-- Velesunio_ambiguus **
 
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