MUSSELpdb | superfamily Hyrioidea Swainson, 1840 cladomics

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Page last updated
Thu 3 September 2025

The Freshwater Mussels (Unionoida) of the World (and other less consequential bivalves)

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Bivalvia | Unionida

superfamily HYRIOIDEA Swainson, 1840: cladomics
1 family

Cladomics Summary

number of species = 93 (124 total)
number of subtaxa (rank: family) = 1
analyzed in 105 cladograms in 55 publications
maximum number of terminals in an analysis = 28 (mean = 4.07, median = 3)
maximum number of subtaxa (family) = 1 (mean = 1, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 61
maximum number of included terminals = 28, (mean = 6.28 median = 5)
maximum number of subtaxa (family) = 1 (mean = 1, median = 1)
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

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

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

tT	bi	riS	oT	roT	roS	iD
28	28	34	1	604	602	2	0

                       +----------------------------- Iridininae                
                       |                                                        
  +--------------------+  +-------------------------- Aspathariinae             
  |                    |  |                                                     
  |                    +--+  +----------------------- Etheria                   
  |                       |  |                                                  
  |                       +--+  +-------------------- Anodontites_trapesialis   
  |                          |  |                                               
  |                          +--+  +----------------- Mulleria                  
  |                             |  |                                            
  |                             +--+  +-------------- Lamproscapha              
  |                                |  |                                         
  |                                +--+           +-- Monocondylaea             
  |                                   |  +--------+                             
  |                                   |  |        +-- Tamsiella                 
  |                                   +--+                                      
  |                                      |  +-------- Mycetopodinae             
--+                                      |  |                                   
  |                                      +--+     +-- Anodontites_guanarensis   
  |                                         |  +--+                             
  |                                         |  |  +-- Fossula                   
  |                                         +--+                                
  |                                            |  +-- Anodontites_trigona       
  |                                            +--+                             
  |                                               +-- Anodontites_elongata      
  |                                                                             
  |  +----------------------------------------------- Hyriidae **                  
  |  |                                                                          
  |  |                          +-------------------- Unioninae                 
  |  |                          |                                               
  |  |  +-----------------------+  +----------------- Lamellidens               
  |  |  |                       |  |                                            
  |  |  |                       +--+              +-- Balwantia                 
  +--+  |                          |  +-----------+                             
     |  |                          |  |           +-- Trapezidens               
     |  |                          +--+                                         
     |  |                             |  +----------- Leoparreysiini            
     |  |                             +--+                                      
     |  |                                |  +-------- Pseudomulleria            
     |  |                                +--+                                   
     +--+                                   |  +----- Indochinellini            
        |                                   +--+                                
        |                                      |  +-- Parreysiini               
        |                                      +--+                             
        |                                         +-- Coelaturini               
        |                                                                       
        |  +----------------------------------------- Rectidentini              
        |  |                                                                    
        |  |  +-------------------------------------- Margaritiferidae          
        |  |  |                                                                 
        +--+  |  +----------------------------------- Ambleminae                
           |  |  |                                                              
           |  |  |     +----------------------------- Indopseudodontina         
           |  |  |     |                                                        
           |  |  |     |                          +-- Lannanaia                 
           +--+  |  +--+  +-----------------------+                             
              |  |  |  |  |                       +-- Pilsbryoconcha            
              |  |  |  |  |                                                     
              |  |  |  +--+  +----------------------- Songkhlanaia              
              |  |  |     |  |                                                  
              |  |  |     |  |                    +-- Nyeinchanconcha           
              |  |  |     +--+  +-----------------+                             
              |  |  |        |  |                 +-- Sundadontina              
              +--+  |        |  |                                               
                 |  |        |  |                 +-- Isannaia                  
                 |  |        +--+              +--+                             
                 |  |           |              |  +-- Namkongnaia               
                 |  |           |  +-----------+                                
                 |  |           |  |           |  +-- Thaiconcha                
                 |  |           |  |           +--+                             
                 |  |           +--+              +-- Bineurus                  
                 |  |              |                                            
                 |  |              |  +-------------- Pseudodon_walpolei        
                 |  |              |  |                                         
                 |  |              +--+  +----------- Pseudodon_vagulus         
                 |  |                 |  |                                      
                 |  |                 |  |     +----- Pseudodon_vondembuschianus
                 +--+                 +--+  +--+                                
                    |                    |  |  |  +-- Pseudodon_mekongi         
                    |                    |  |  +--+                             
                    |                    +--+     +-- Schepmaniini              
                    |                       |                                   
                    |                       |  +----- Pseudodon_lenyanensis__2  
                    |                       +--+                                
                    |                          |  +-- Pseudodon_inoscularis     
                    |                          +--+                             
                    |                             +-- Pseudodon_lenyanensis__1  
                    |                                                           
                    |           +-------------------- Chamberlainiini           
                    |           |                                               
                    |        +--+  +----------------- Parvasolenaia             
                    |        |  |  |                                            
                    |        |  +--+  +-------------- Pseudopostolata           
                    |        |     |  |                                         
                    |        |     +--+  +----------- Koreosolenaia             
                    |        |        |  |                                      
                    |        |        +--+        +-- Cosmopseudodon            
                    |        |           |  +-----+                             
                    |        |           |  |     +-- Guiunio                   
                    |        |           +--+                                   
                    +--------+              |  +----- Ptychorhynchus            
                             |              +--+                                
                             |                 |  +-- Obovalis                  
                             |                 +--+                             
                             |                    +-- Postolata                 
                             |                                                  
                             |                    +-- Caudiculatus              
                             |     +--------------+                             
                             |     |              +-- Lamprotulini              
                             |     |                                            
                             +-----+  +-------------- Sinosolenaia_carinata     
                                   |  |                                         
                                   |  |        +----- Gonidea                   
                                   +--+  +-----+                                
                                      |  |     |  +-- Microcondylaea            
                                      |  |     +--+                             
                                      +--+        +-- Leguminaia                
                                         |                                      
                                         |  +-------- Inversidens               
                                         +--+                                   
                                            |  +----- Sinosolenaia_recognita    
                                            +--+                                
                                               |  +-- Sinosolenaia_iridinea     
                                               +--+                             
                                                  +-- Sinosolenaia_oleivora

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

tT	bi	riS	oT	roT	roS	iD
19	19	20	1	1	0	0	22.5

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

tT	bi	riS	oT	roT	roS	iD
19	19	21	1	1	0	0	25.5

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

tT	bi	riS	oT	roT	roS	iD
14	14	12	1	6	5	2	37

  +----- Hyriidae **   
--+                 
  |  +-- Unionoidea 
  +--+              
     +-- Etherioidea

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

tT	bi	riS	oT	roT	roS	iD
14	14	12	1	185	183	2	41

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
14	14	12	1	187	185	2	39

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
14	14	9	1	187	185	2	38

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
10	10	7	1	26	25	2	29

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
10	10	8	1	4	4	1	31.5

     +-- Margaritiferidae
  +--+                   
  |  +-- Unioninae       
--+                      
  |  +-- Ambleminae      
  +--+                   
     +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
9	9	7	1	38	37	2	32.5

  +----- Unionoidea 
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Etherioidea

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

tT	bi	riS	oT	roT	roS	iD
9	9	7	1	37	36	2	32.5

  +----------------- Parreysiinae    
  |                                  
  |           +----- Margaritiferidae
--+  +--------+                      
  |  |        |  +-- Hyriidae **        
  |  |        +--+                   
  +--+           +-- Etherioidea     
     |                               
     |  +----------- Pseudodontini   
     +--+                            
        |  +-------- Rectidentini    
        +--+                         
           |  +----- Unioninae       
           +--+                      
              |  +-- Gonideini       
              +--+                   
                 +-- Ambleminae

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

tT	bi	riS	oT	roT	roS	iD
9	9	7	1	21	18	2	38

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
9	9	7	1	130	128	2	38

  +-------- Margaritiferidae
--+                         
  |  +----- Unionidae       
  +--+                      
     |  +-- Etherioidea     
     +--+                   
        +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
9	9	7	1	21	18	2	38

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
9	9	7	1	38	37	2	32.5

  +-------------------------- Hyriidae **        
  |                                           
  |                 +-------- Iridinidae      
--+  +--------------+                         
  |  |              |  +----- Etheria         
  |  |              +--+                      
  |  |                 |  +-- Mulleria        
  +--+                 +--+                   
     |                    +-- Mycetopodidae   
     |                                        
     |  +-------------------- Margaritiferidae
     +--+                                     
        |  +----------------- Parreysiinae    
        +--+                                  
           |  +-------------- Pseudomulleria  
           +--+                               
              |  +----------- Unioninae       
              +--+                            
                 |  +-------- Rectidentini    
                 +--+                         
                    |  +----- Ambleminae      
                    +--+                      
                       |  +-- Gonideini       
                       +--+                   
                          +-- Pseudodontini

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

tT	bi	riS	oT	roT	roS	iD
9	9	7	1	37	36	2	32.5

  +-------------------------- Hyriidae **        
  |                                           
  |                       +-- Etheria         
  |                    +--+                   
--+                    |  +-- Mulleria        
  |  +-----------------+                      
  |  |                 |  +-- Mycetopodidae   
  |  |                 +--+                   
  +--+                    +-- Iridinidae      
     |                                        
     |  +-------------------- Parreysiinae    
     +--+                                     
        |  +----------------- Pseudomulleria  
        +--+                                  
           |  +-------------- Margaritiferidae
           +--+                               
              |  +----------- Pseudodontini   
              +--+                            
                 |  +-------- Rectidentini    
                 +--+                         
                    |  +----- Unioninae       
                    +--+                      
                       |  +-- Gonideini       
                       +--+                   
                          +-- Ambleminae

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

tT	bi	riS	oT	roT	roS	iD
9	9	7	1	130	128	2	38

                       +-- Chamberlainia   
  +--------------------+                   
  |                    +-- Margaritiferidae
  |                                        
  |                    +-- Etherioidea     
--+--------------------+                   
  |                    +-- Hyriidae **        
  |                                        
  |  +-------------------- Parreysiinae    
  +--+                                     
     |  +----------------- Unioninae       
     +--+                                  
        |  +-------------- Rectidentini    
        +--+                               
           |  +----------- Ambleminae      
           +--+                            
              |  +-------- Pseudodontini   
              +--+                         
                 |  +----- Gonideini       
                 +--+                      
                    |  +-- Sinohyriopsis   
                    +--+                   
                       +-- Lamprotulini

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

tT	bi	riS	oT	roT	roS	iD
8	8	9	1	1	1	1	35

  +-- Unionoidea
--+             
  +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	1	1	1	1	37

  +-- Unionoidea
--+             
  +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	6	1	0	0	0	29

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

tT	bi	riS	oT	roT	roS	iD
6	6	5	1	5	5	1	36.5

  +-- Margaritiferidae
  |                   
--+-- Unionidae       
  |                   
  +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	1	26	25	2	33.5

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	1	36	35	2	35

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	1	26	25	2	33

  +-------- Margaritiferidae
--+                         
  |  +----- Unionidae       
  +--+                      
     |  +-- Etherioidea     
     +--+                   
        +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	1	135	134	2	33

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	1	36	35	2	33

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	1	0	0	0	31

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	1	29	28	2	33

  +-------------------- Hyriidae **        
  |                                     
  |           +-------- Iridinidae      
--+  +--------+                         
  |  |        |  +----- Etheria         
  |  |        +--+                      
  |  |           |  +-- Mulleria        
  +--+           +--+                   
     |              +-- Mycetopodidae   
     |                                  
     |  +-------------- Parreysiinae    
     |  |                               
     +--+  +----------- Pseudomulleria  
        |  |                            
        +--+  +-------- Anodontini      
           |  |                         
           +--+  +----- Gonideinae      
              |  |                      
              +--+----- Unionini        
                 |                      
                 |  +-- Margaritiferidae
                 +--+                   
                    +-- Ambleminae

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	1	26	25	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
5	5	2	1	26	25	2	36

  +----- Unionoidea 
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
5	5	1	1	26	25	2	35

  +----------------- Margaritiferidae
  |                                  
  |  +-------------- Amblemini       
  |  |                               
  |  +-------------- Parreysiinae    
  |  |                               
  |  +-------------- Eurynia         
--+  |                               
  |  +-------------- Fusconaia       
  |  |                               
  |  +-------------- Gonideinae      
  |  |                               
  |  +-------------- Pleurobema      
  |  |                               
  +--+-------------- Quadrulini      
     |                               
     |  +----------- Ortmanniana     
     +--+                            
     |  |  +-------- Glebula         
     |  +--+                         
     |     |  +----- Potamilus       
     |     +--+                      
     |        |  +-- Ligumia         
     |        +--+                   
     |           +-- Toxolasma       
     |                               
     |  +----------- Cyrtonaias      
     +--+                            
        |  +-------- Unionini        
        +--+                         
           |  +----- Anodontini      
           +--+                      
              |  +-- Hyriidae **        
              +--+                   
                 +-- Etherioidea

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

tT	bi	riS	oT	roT	roS	iD
4	4	2	1	0	0	0	34

Graf, 2000 [Publ 2647], fig. 1

tT	bi	riS	oT	roT	roS	iD
4	4	2	1	13	12	2	34.5

  +-------------- Margaritiferidae
  |                               
  |     +-------- Unionini        
--+  +--+                         
  |  |  |  +----- Pleurobemini    
  |  |  +--+                      
  |  |     |  +-- Anodontini      
  +--+     +--+                   
     |        +-- Lampsilini      
     |                            
     |  +-------- Parreysiini     
     +--+                         
        |  +----- Coelaturini     
        +--+                      
           |  +-- Hyriidae **        
           +--+                   
              +-- Etherioidea

Graf, 2002 [Publ 2651], fig. 2

tT	bi	riS	oT	roT	roS	iD
4	4	2	1	14	13	1	31.5

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
4	4	2	1	32	31	2	34

  +-------- Hyriidae **        
--+                         
  |  +----- Margaritiferidae
  +--+                      
     |  +-- Etherioidea     
     +--+                   
        +-- Unionidae

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

tT	bi	riS	oT	roT	roS	iD
4	4	2	1	2	2	1	32

  +-- Unionoidea
--+             
  +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
4	4	2	1	32	31	2	34

  +-------- Etherioidea     
--+                         
  |  +----- Unionidae       
  +--+                      
     |  +-- Margaritiferidae
     +--+                   
        +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
4	4	2	1	36	35	2	34

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	12	10	2	33

  +-------- Hyriidae **        
--+                         
  |  +----- Unionidae       
  +--+                      
     |  +-- Margaritiferidae
     +--+                   
        +-- Etherioidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	12	10	2	35

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	25	23	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	0	1	12	9	1	34

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	31	29	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	0	1	12	9	1	34

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	66	64	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	66	64	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	66	64	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	13	10	1	33

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	31	29	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	88	86	2	35

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	88	86	2	35

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	0	0	0	33

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	28	26	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	26	24	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	1	25	23	2	33

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	1	120	118	2	--

  +-------- Hyriidae **        
--+                         
  |  +----- Margaritiferidae
  +--+                      
     |  +-- Etherioidea     
     +--+                   
        +-- Unionidae

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	1	24	22	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	1	79	78	2	--

  +-------- Hyriidae **        
--+                         
  |  +----- Margaritiferidae
  +--+                      
     |  +-- Etherioidea     
     +--+                   
        +-- Unionidae

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	1	45	44	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	1	20	20	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	1	20	20	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	4	3	2	--

  +-------- Hyriidae **        
--+                         
  |  +----- Unionidae       
  +--+                      
     |  +-- Etherioidea     
     +--+                   
        +-- Margaritiferidae

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	72	71	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	37	37	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	24	23	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	14	13	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	13	13	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	56	55	2	--

  +-------- Etherioidea     
--+                         
  |  +----- Unionidae       
  +--+                      
     |  +-- Hyriidae **        
     +--+                   
        +-- Margaritiferidae

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	10	10	1	--

  +----- Unionidae       
--+                      
  |  +-- Margaritiferidae
  +--+                   
     +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	14	13	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	14	13	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	22	22	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	84	84	2	--

  +-------- Etherioidea     
--+                         
  |  +----- Unionidae       
  +--+                      
     |  +-- Hyriidae **        
     +--+                   
        +-- Margaritiferidae

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	72	71	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	17	17	1	--

  +----------- Cristariina    
  |                           
  |  +-------- Lanceolariini  
  |  |                        
--+  +-------- Anodonta       
  |  |                        
  |  +-------- Unionini       
  |  |                        
  +--+     +-- Alasmidontina  
     +-----+                  
     |     +-- Pseudanodonta  
     |                        
     |  +----- Ambleminae     
     |  |                     
     +--+  +-- Hyriidae **       
        |  |                  
        +--+-- Lamprotulini   
           |                  
           +-- Chamberlainiini

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	45	45	2	--

  +-------- Unionidae       
--+                         
  |  +----- Margaritiferidae
  +--+                      
     |  +-- Hyriidae **        
     +--+                   
        +-- Etherioidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	90	89	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	4	3	2	--

     +-- Etherioidea     
  +--+                   
  |  +-- Margaritiferidae
--+                      
  |  +-- Unionidae       
  +--+                   
     +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	72	72	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	7	7	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	47	46	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	90	89	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	5	3	2	--

  +----- Unionidae       
  |                      
--+----- Hyriidae **        
  |                      
  |  +-- Etherioidea     
  +--+                   
     +-- Margaritiferidae

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	29	28	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	55	54	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	94	93	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	29	28	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	38	38	1	--

  +-------------- Margaritiferidae
  |                               
--+  +----------- Unioninae       
  |  |                            
  +--+  +-------- Lampsilini      
     |  |                         
     +--+     +-- Hyriidae **        
        |  +--+                   
        |  |  +-- Amblemini       
        +--+                      
           |  +-- Pleurobemini    
           +--+                   
              +-- Quadrulini

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	94	93	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	5	3	2	--

     +-- Etherioidea     
  +--+                   
  |  +-- Margaritiferidae
--+                      
  |  +-- Unionidae       
  +--+                   
     +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	67	66	2	--

  +----- Hyriidae **   
--+                 
  |  +-- Etherioidea
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	16	16	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	8	8	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	24	23	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	5	3	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	10	10	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	41	40	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	5	3	2	--

  +-------- Hyriidae **        
--+                         
  |  +----- Unionidae       
  +--+                      
     |  +-- Etherioidea     
     +--+                   
        +-- Margaritiferidae

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	56	55	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	30	30	1	--

  +-- Margaritiferidae
  |                   
--+-- Hyriidae **        
  |                   
  +-- Unionidae

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	2	2	1	--

  +-- Etheriidae
  |             
--+-- Iridinidae
  |             
  +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	9	9	1	--

  +----- Unionidae       
--+                      
  |  +-- Margaritiferidae
  +--+                   
     +-- Hyriidae **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	7	7	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	86	86	2	--

  +----- Etherioidea
--+                 
  |  +-- Hyriidae **   
  +--+              
     +-- Unionoidea

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	8	8	1	--

  +-- Hyriidae **  
--+             
  +-- Unionoidea

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