Bivalvia | Unionida | Unionoidea

family MARGARITIFERIDAE Henderson, 1929: cladomics

6 genera, 2 subfamilies

Cladomics pages are experimental — even more so than the rest of this site!

The purpose of the cladomics pages for each genus and family-group level taxon is to provide statistics and references indicating the extent to which published phylogenetic work supports the classification used here.

The Cladomics Summary provides an overview of the number of studies that have included terminal species representing this taxon as well as an indication of how frequently it has been recovered as monophyletic.

Following the summary, the available published clades representing the taxon are listed.

For more information, visit the Cladomics FARQ (frequently asked rhetorical questions) page. Abbreviations for the various statistics are explained on the FARQ!

Cladomics Summary

number of species = 16 (46 total)
number of subtaxa (rank: subfamily) = 8
analyzed in 269 cladograms in 152 publications
maximum number of terminals in an analysis = 16 (mean = 3.2, median = 2)
maximum number of subtaxa (subfamily) = 2 (mean = 1.28, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 184
maximum number of included terminals = 16, (mean = 4.21 median = 2)
maximum number of subtaxa (subfamily) = 2 (mean = 1.41, median = 1)
proportion of cladograms that support basic monophyly = 0.96

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 74
(Rigorous monophyly refers to cladograms with terminal species representing more than one subtaxon (subfamily) taxon and at least one rigorous outgroup (family).
maximum number of included terminals = 15 (mean = 6.07, median = 5.5)
maximum number of subtaxa (subfamily) = 2 (mean = 2, median = 2)
proportion of cladograms that support rigorous monophyly = 0.97


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 of Margaritiferidae cladogram distances
PCoA Ordination

NJ Tree of Margaritiferidae cladogram distances
NJ Tree


Cladograms

Zhou et al., 2007 [Publ 2658], fig. 1

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
3 21 19 2 18 18 1 0
  +-------------------- Margaritiferinae **       
  |                                            
  |  +----------------- Anodontini             
--+  |                                         
  |  |     +----------- Chamberlainiini        
  |  |  +--+                                   
  +--+  |  |  +-------- Lamprotula_leaii       
     |  |  +--+                                
     |  |     |  +----- Lamprotula_caveata     
     |  |     +--+                             
     |  |        |  +-- Gibbosulinae **           
     +--+        +--+                          
        |           +-- Lamprotula_cornuumlunae
        |                                      
        |        +----- Aculamprotulina        
        |     +--+                             
        |     |  |  +-- Tchangsinaia           
        |     |  +--+                          
        +-----+     +-- Nodularia              
              |                                
              |  +----- Schistodesmus          
              +--+                             
                 |  +-- Lanceolariini          
                 +--+                          
                    +-- Acuticostini           

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

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
14 14 12 2 617 567 1 29
  +----------------------------------------- Rectidentini              
  |                                                                    
  |                                +-------- Pseudunio **                 
  |  +-----------------------------+                                   
  |  |                             |  +----- Gibbosulinae **              
  |  |                             +--+                                
--+  |                                |  +-- Cumberlandia **              
  |  |                                +--+                             
  |  |                                   +-- Margaritifera **             
  |  |                                                                 
  |  |  +----------------------------------- 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     

Lyubas et al., 2019 [Publ 3963], fig. 10

iTtTbiriSoTroTroSiD
16 16 12 2 0 0 0 31
  +-- Gibbosulinae **    
--+                   
  +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
15 15 13 2 11 2 1 30
  +-------------- Unionidae                
  |                                        
--+  +----------- Pseudunio_marocanus **      
  |  |                                     
  |  |        +-- Pseudunio_auricularius__1 **
  +--+        |                            
     |  +-----+-- Pseudunio_auricularius__2 **
     |  |     |                            
     +--+     +-- Pseudunio_homsensis **      
        |                                  
        |  +----- Gibbosulinae **             
        +--+                               
           |  +-- Cumberlandia **             
           +--+                            
              +-- Margaritifera **            

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

iTtTbiriSoTroTroSiD
14 14 12 2 127 114 1 29
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
14 14 12 2 14 3 1 29
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
14 14 12 2 14 3 1 29
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
13 13 9 2 187 161 1 28
  +-------- Unionidae               
--+                                 
  |  +----- Gibbosula_rochechouartii **
  +--+                              
     |  +-- Gibbosula_laosensis **     
     +--+                           
        +-- Margaritiferinae **        

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

iTtTbiriSoTroTroSiD
13 13 11 2 185 159 1 28
  +-------- Unionidae               
--+                                 
  |  +----- Gibbosula_rochechouartii **
  +--+                              
     |  +-- Gibbosula_laosensis **     
     +--+                           
        +-- Margaritiferinae **        

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

iTtTbiriSoTroTroSiD
13 13 9 2 187 161 1 28
  +-------- Unionidae               
--+                                 
  |  +----- Gibbosula_rochechouartii **
  +--+                              
     |  +-- Gibbosula_laosensis **     
     +--+                           
        +-- Margaritiferinae **        

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

iTtTbiriSoTroTroSiD
12 12 10 2 56 45 1 29
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Pseudunio **    
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
12 12 10 2 22 11 1 29
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Pseudunio **    

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

iTtTbiriSoTroTroSiD
12 12 10 2 22 11 1 27
  +-------- Unionidae               
--+                                 
  |  +----- Gibbosula_rochechouartii **
  +--+                              
     |  +-- Gibbosula_laosensis **     
     +--+                           
        +-- Margaritiferinae **        

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

iTtTbiriSoTroTroSiD
12 12 10 2 56 45 1 29
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Pseudunio **    

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

iTtTbiriSoTroTroSiD
12 12 10 2 17 6 1 27
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
12 12 10 2 56 45 1 29
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Pseudunio **    
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Margaritifera **

Araujo et al., 2016 [Publ 3746], fig. 1_2b

iTtTbiriSoTroTroSiD
11 11 9 2 2 1 1 27
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Vikhrev et al., 2018 [Publ 3812], fig. 3

iTtTbiriSoTroTroSiD
11 11 9 2 2 2 1 27
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Araujo et al., 2016 [Publ 3746], fig. 2a

iTtTbiriSoTroTroSiD
11 11 9 2 0 0 0 29
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae ** 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
10 10 8 2 66 66 1 28
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Pseudunio **    

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

iTtTbiriSoTroTroSiD
10 10 7 2 128 113 1 28
  +-------- Unionidae    
  |                      
--+  +----- Cumberlandia ** 
  |  |                   
  +--+  +-- Gibbosulinae ** 
     |  |                
     +--+-- Pseudunio **    
        |                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
10 10 8 2 20 4 1 28
  +----------- Unionidae    
--+                         
  |  +-------- Pseudunio **    
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Cumberlandia ** 

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

iTtTbiriSoTroTroSiD
10 10 8 2 66 66 1 28
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Gibbosulinae ** 
     +--+                   
        |  +-- Pseudunio **    
        +--+                
           +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
10 10 8 2 20 4 1 28
  +----------- Unionidae    
--+                         
  |  +-------- Pseudunio **    
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Cumberlandia ** 

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

iTtTbiriSoTroTroSiD
10 10 8 2 128 113 1 28
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae ** 
     +--+                
        +-- Pseudunio **    

Bolotov et al., 2015 [Publ 3646], fig. 2A

iTtTbiriSoTroTroSiD
8 8 5 1 1 1 1 25
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Bolotov et al., 2015 [Publ 3646], fig. 2B

iTtTbiriSoTroTroSiD
8 8 6 1 1 1 1 25
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Inoue et al., 2013 [Publ 3579], fig. 3

iTtTbiriSoTroTroSiD
8 8 6 1 1 1 1 25
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Takeuchi et al., 2015 [Publ 3661], fig. 2

iTtTbiriSoTroTroSiD
8 8 6 1 2 1 1 25
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Takeuchi et al., 2015 [Publ 3661], fig. 3

iTtTbiriSoTroTroSiD
8 8 6 1 2 1 1 25
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 89 85 1 22
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huff et al., 2004 [Publ 2697], fig. 4

iTtTbiriSoTroTroSiD
7 7 4 1 4 3 1 24
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Huff et al., 2004 [Publ 2697], fig. 5-6

iTtTbiriSoTroTroSiD
7 7 5 1 5 3 1 24
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 67 63 1 22
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 34 21 1 23
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 89 85 1 22
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 84 74 1 23
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 67 63 1 22
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Araujo et al., 2009 [Publ 2395], fig. 3

iTtTbiriSoTroTroSiD
7 7 5 1 1 1 1 24
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Gomes-dos-Santos et al., 2019 [Publ 3943], fig. 1

iTtTbiriSoTroTroSiD
7 7 5 2 3 3 1 22
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huff et al., 2004 [Publ 2697], fig. 2

iTtTbiriSoTroTroSiD
7 7 3 1 5 3 1 25
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 62 58 1 22
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 34 21 1 23
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
7 7 5 2 84 74 1 23
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
6 6 4 1 30 10 1 23
  +-------- Unioninae       
--+                         
  |  +----- Margaritiferinae **
  +--+                      
     |  +-- Gonideinae      
     +--+                   
        +-- Ambleminae      

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

iTtTbiriSoTroTroSiD
6 6 4 2 68 65 1 21
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
6 6 4 2 50 46 1 21
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huff et al., 2004 [Publ 2697], fig. 3

iTtTbiriSoTroTroSiD
6 6 2 1 2 1 1 23
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
6 6 4 2 36 32 1 21
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
6 6 4 2 82 79 1 21
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
6 6 4 2 80 77 1 21
  +-- Gibbosulinae **    
--+                   
  +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
5 5 0 2 112 112 1 23
  +----- Unionidae                  
  |                                 
  |  +-- Gibbosulinae **               
--+  |                              
  |  +-- Margaritifera_dahurica **     
  |  |                              
  +--+-- Margaritifera_margaritifera **
     |                              
     +-- Margaritifera_laevis **       
     |                              
     +-- Margaritifera_middendorffi ** 

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

iTtTbiriSoTroTroSiD
4 4 2 2 42 40 1 20
  +-- Gibbosulinae **    
--+                   
  +-- Margaritiferinae **

Bolotov et al., 2020 [Publ 4084], fig. 1

iTtTbiriSoTroTroSiD
4 4 0 1 32 32 1 23
  +----- Margaritiferinae **
  |                      
--+----- Ambleminae      
  |                      
  |  +-- Gonideinae      
  +--+                   
     +-- Unioninae       

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

iTtTbiriSoTroTroSiD
4 4 2 2 45 41 1 20
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
4 4 2 1 35 34 1 21
1

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

iTtTbiriSoTroTroSiD
4 4 2 1 34 33 1 21
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Rosenberg et al., 1994 [Publ 2719], fig. 3

iTtTbiriSoTroTroSiD
3 19 5 1 16 16 1 36
  +-- Cumberlandia ** 
  |                
--+-- Margaritifera **
  |                
  +-- Unionidae    

Froufe et al., 2019 [Publ 3936], fig. 3

iTtTbiriSoTroTroSiD
3 3 1 1 41 41 1 23
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
3 3 1 1 19 17 1 20
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Gomes-dos-Santos et al., 2019 [Publ 3943], fig. 1-M

iTtTbiriSoTroTroSiD
3 3 1 1 3 3 1 20
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Mock et al., 2013 [Publ 3598], fig. 3

iTtTbiriSoTroTroSiD
3 3 1 1 0 0 0 22
1

Froufe et al., 2019 [Publ 3936], fig. 6

iTtTbiriSoTroTroSiD
3 3 1 1 41 41 1 23
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Machordom et al., 2003 [Publ 2688], fig. 3

iTtTbiriSoTroTroSiD
3 3 1 1 1 1 1 20
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Fukata & Iigo, 2020 [Publ 4106], fig. 1

iTtTbiriSoTroTroSiD
3 3 1 2 51 51 1 21
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lydeard et al., 1996 [Publ 2656], fig. 3

iTtTbiriSoTroTroSiD
3 3 1 1 24 24 1 20
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Cha et al., 2018 [Publ 3916], fig. 1

iTtTbiriSoTroTroSiD
3 3 1 1 35 35 1 23
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Rosenberg et al., 1997 [Publ 2720], fig. 2A

iTtTbiriSoTroTroSiD
3 22 4 1 19 18 1 37
  +----- Cumberlandia ** 
  |                   
  +----- None__2      
--+                   
  +----- Margaritifera **
  |                   
  |  +-- None__1      
  +--+                
     +-- Unionidae    

Aunins et al., 2018 [Publ 3863], fig. 1

iTtTbiriSoTroTroSiD
3 3 1 1 37 37 1 21
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
3 3 1 1 19 17 1 20
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2017 [Publ 3792], fig. 2

iTtTbiriSoTroTroSiD
3 3 1 1 29 29 1 23
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
3 3 1 1 44 41 1 20
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Rosenberg et al., 1997 [Publ 2720], fig. 2B

iTtTbiriSoTroTroSiD
3 22 3 1 19 18 1 36
  +-- Cumberlandia ** 
  |                
  +-- None__1      
  |                
--+-- None__2      
  |                
  +-- Margaritifera **
  |                
  +-- Unionidae    

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

iTtTbiriSoTroTroSiD
3 3 1 1 11 10 1 20
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Sano et al., 2020 [Publ 4138], fig. 1

iTtTbiriSoTroTroSiD
3 3 1 2 31 30 1 21
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2020 [Publ 4147], fig. 4

iTtTbiriSoTroTroSiD
3 3 1 2 11 11 1 20
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Stewart et al., 2021 [Publ 4451], fig. 1

iTtTbiriSoTroTroSiD
3 3 1 1 13 13 1 22
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
3 3 1 1 45 45 1 23
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lydeard et al., 1996 [Publ 2656], fig. 4

iTtTbiriSoTroTroSiD
3 3 1 1 25 24 1 20
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Dai et al., 2024 [Publ 4386], fig. 5-BI

iTtTbiriSoTroTroSiD
2 2 0 2 20 20 1 --
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **    

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 29 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Wu et al., 2023 [Publ 4339], fig. 5

iTtTbiriSoTroTroSiD
2 2 0 1 17 17 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 2 45 45 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2018 [Publ 3836], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 1 51 51 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 24 1 --
1

Chen et al., 2025 [Publ 4487], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 2 39 39 1 --
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **    

Lopes-Lima et al., 2020 [Publ 4080], fig. 12

iTtTbiriSoTroTroSiD
2 2 0 1 6 6 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Chen et al., 2023 [Publ 4302], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 2 39 39 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
2 2 0 2 44 44 1 --
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **    

Bolotov et al., 2016 [Publ 3732], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 2 6 6 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Amaro et al., 2017 [Publ 3791], fig. 3A

iTtTbiriSoTroTroSiD
2 2 0 1 10 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 28 25 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Vikhrev et al., 2017 [Publ 3810], fig. 4

iTtTbiriSoTroTroSiD
2 2 0 2 8 8 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2020 [Publ 4080], fig. 7

iTtTbiriSoTroTroSiD
2 2 0 1 6 6 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 25 25 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 90 86 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Chen et al., 2025 [Publ 4487], fig. 3

iTtTbiriSoTroTroSiD
2 2 0 2 39 39 1 --
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **    

Huang et al., 2002 [Publ 2657], fig. 1

iTtTbiriSoTroTroSiD
2 34 0 1 32 32 1 40
  +-- Cumberlandia ** 
  |                
--+-- Margaritifera **
  |                
  +-- Unionidae    

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

iTtTbiriSoTroTroSiD
2 2 0 2 93 93 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2020 [Publ 4080], fig. 5

iTtTbiriSoTroTroSiD
2 2 0 1 7 7 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 24 1 --
1

Sano et al., 2017 [Publ 3799], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 1 19 17 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Bespalaya et al., 2018 [Publ 3811], fig. 3

iTtTbiriSoTroTroSiD
2 2 0 2 8 8 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Teiga-Teixeira et al., 2020 [Publ 4173], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 1 17 17 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Pavan-Kumar et al., 2022 [Publ 4263], fig. S5

iTtTbiriSoTroTroSiD
2 2 0 1 37 37 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Dai et al., 2026 [Publ 4757], fig. 3

iTtTbiriSoTroTroSiD
2 2 0 2 32 32 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2020 [Publ 4080], fig. 17

iTtTbiriSoTroTroSiD
2 2 0 1 4 4 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2019 [Publ 3901], fig. 4

iTtTbiriSoTroTroSiD
2 2 0 1 24 24 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 2 46 46 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Amaro et al., 2017 [Publ 3791], fig. 3B

iTtTbiriSoTroTroSiD
2 2 0 1 10 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Burzynski & Soroka, 2018 [Publ 3862], fig. 5a

iTtTbiriSoTroTroSiD
2 2 0 1 18 18 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 120 111 1 --
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 34 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 45 25 1 --
1

Lopes-Lima et al., 2020 [Publ 4080], fig. 14

iTtTbiriSoTroTroSiD
2 2 0 1 4 4 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Feng et al., 2016 [Publ 3800], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 1 12 12 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Campbell et al., 2005 [Publ 2660], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 1 100 100 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2020 [Publ 4080], fig. 10

iTtTbiriSoTroTroSiD
2 2 0 1 12 12 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Bolotov et al., 2022 [Publ 4250], fig. 8

iTtTbiriSoTroTroSiD
2 7 0 2 47 47 1 25
     +----- Parreysiinae    
  +--+                      
  |  |  +-- Pseudodontini   
  |  +--+                   
  |     +-- Rectidentini    
  |                         
--+     +-- Gibbosulinae **    
  |     |                   
  |     +-- Margaritiferinae **
  |     |                   
  |     +-- Sinosolenaia    
  |     |                   
  +-----+-- Leguminaia      
        |                   
        +-- Gonidea         
        |                   
        +-- Lamprotula      
        |                   
        +-- Potomida        

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

iTtTbiriSoTroTroSiD
2 2 0 2 43 43 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2020 [Publ 4080], fig. 9

iTtTbiriSoTroTroSiD
2 2 0 1 4 4 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 43 43 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Dai et al., 2025 [Publ 4440], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 2 43 43 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Machordom et al., 2003 [Publ 2688], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 1 1 1 1 --
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
2 2 0 1 23 19 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Pavan-Kumar et al., 2022 [Publ 4263], fig. 4

iTtTbiriSoTroTroSiD
2 2 0 1 35 35 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 18 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2020 [Publ 4080], fig. 8

iTtTbiriSoTroTroSiD
2 2 0 1 5 5 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Dai et al., 2024 [Publ 4386], fig. 5-ML

iTtTbiriSoTroTroSiD
2 2 0 2 20 20 1 --
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **    

Huang et al., 2019 [Publ 3843], fig. S1

iTtTbiriSoTroTroSiD
2 2 0 2 45 45 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
2 2 0 2 43 43 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Dai et al., 2024 [Publ 4385], fig. 3-BI

iTtTbiriSoTroTroSiD
2 2 0 2 29 29 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Chen et al., 2023 [Publ 4302], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 2 39 39 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
2 2 0 1 38 30 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 18 1 --
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 18 1 --
  +----------- Parreysiinae    
--+                            
  |  +-------- Margaritiferinae **
  +--+                         
     |  +----- Gonideinae      
     +--+                      
        |  +-- Unioninae       
        +--+                   
           +-- Ambleminae      

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

iTtTbiriSoTroTroSiD
2 2 0 1 44 24 1 --
1

Välilä et al., 2017 [Publ 3793], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 1 1 1 1 --
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 18 1 --
1

Hou et al., 2024 [Publ 4400], fig. 4

iTtTbiriSoTroTroSiD
2 2 0 1 74 74 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 90 86 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 34 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 55 50 1 --
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 51 51 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Dai et al., 2024 [Publ 4385], fig. 3-ML

iTtTbiriSoTroTroSiD
2 2 0 2 29 29 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2026 [Publ 4811], fig. 3

iTtTbiriSoTroTroSiD
2 2 0 2 15 15 1 --
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **    

Lopes-Lima et al., 2019 [Publ 3901], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 1 26 26 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Dai et al., 2025 [Publ 4440], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 2 43 43 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 21 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Krebs et al., 2003 [Publ 2691], fig. 3

iTtTbiriSoTroTroSiD
2 2 0 1 31 31 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Huang et al., 2002 [Publ 2657], fig. 2

iTtTbiriSoTroTroSiD
2 34 0 1 32 32 1 39
  +-- Cumberlandia ** 
  |                
--+-- Margaritifera **
  |                
  +-- Unionidae    

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

iTtTbiriSoTroTroSiD
2 2 0 1 51 51 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 45 25 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 78 69 1 --
1

Wu et al., 2024 [Publ 4379], fig. 5

iTtTbiriSoTroTroSiD
2 2 0 2 52 52 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 34 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Bogan et al., 2023 [Publ 4295], fig. 3

iTtTbiriSoTroTroSiD
2 2 0 2 29 29 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Sano et al., 2017 [Publ 3799], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 1 31 29 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2020 [Publ 4080], fig. 11

iTtTbiriSoTroTroSiD
2 2 0 1 10 10 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Riccardi et al., 2020 [Publ 4089], fig. 2

iTtTbiriSoTroTroSiD
2 2 0 1 17 17 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2020 [Publ 4080], fig. 16

iTtTbiriSoTroTroSiD
2 2 0 1 6 6 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2019 [Publ 3901], fig. 3

iTtTbiriSoTroTroSiD
2 2 0 1 27 27 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 2 49 49 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wen et al., 2017 [Publ 3795], fig. 3

iTtTbiriSoTroTroSiD
2 2 0 1 14 14 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Wu et al., 2024 [Publ 4379], fig. 4_6

iTtTbiriSoTroTroSiD
2 2 0 1 21 21 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2017 [Publ 3734], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 1 15 14 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Campbell et al., 2005 [Publ 2660], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 1 100 100 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2020 [Publ 4080], fig. 13

iTtTbiriSoTroTroSiD
2 2 0 1 8 8 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 18 1 --
  +----------- Parreysiinae    
--+                            
  |  +-------- Margaritiferinae **
  +--+                         
     |  +----- Ambleminae      
     +--+                      
        |  +-- Gonideinae      
        +--+                   
           +-- Unioninae       

Dai et al., 2024 [Publ 4358], fig. 4

iTtTbiriSoTroTroSiD
2 2 0 2 51 51 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wang et al., 2025 [Publ 4488], fig. 3B

iTtTbiriSoTroTroSiD
2 2 0 2 63 63 1 --
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 34 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 23 19 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 28 25 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Amaro et al., 2017 [Publ 3791], fig. 2

iTtTbiriSoTroTroSiD
2 15 0 1 14 13 1 33
  +-------- Margaritifera_margaritifera__2 **
--+                                       
  |  +----- Unionidae                     
  +--+                                    
     |  +-- Margaritifera_margaritifera__1 **
     +--+                                 
        +-- Margaritifera_falcata **         

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 34 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 21 20 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 44 24 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Huang et al., 2019 [Publ 3843], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 2 43 43 1 --
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **    

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

iTtTbiriSoTroTroSiD
2 2 0 1 55 50 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lopes-Lima et al., 2020 [Publ 4080], fig. 6

iTtTbiriSoTroTroSiD
2 2 0 1 9 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Fonseca et al., 2016 [Publ 3775], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 1 29 29 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 34 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
2 2 0 1 32 18 1 --
  +-------- Anodontini      
  |                         
--+  +----- Gonideinae      
  |  |                      
  +--+----- Unionini        
     |                      
     |  +-- Margaritiferinae **
     +--+                   
        +-- Ambleminae      

Zanatta & Murphy, 2006 [Publ 2672], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 56 56 1 --
  +-- Unioninae       
  |                   
--+-- Margaritiferinae **
  |                   
  +-- Ambleminae      

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 22 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Bashê, 2022 [Publ 4265], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 5 5 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 4 1 --
-- Cumberlandia_monodonta **

Inoue et al., 2018 [Publ 3858], fig. 1_3a-c

iTtTbiriSoTroTroSiD
1 1 0 1 104 104 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Hoeh et al., 1998 [Publ 2695], fig. 4

iTtTbiriSoTroTroSiD
1 1 0 1 4 2 1 --
  +----- Ambleminae      
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Unioninae       

Douda et al., 2024 [Publ 4364], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 13 13 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Nagel et al., 1998 [Publ 1946], fig. 9

iTtTbiriSoTroTroSiD
1 1 0 1 5 5 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 4 1 1 --
-- Margaritifera_margaritifera **

Curole & Kocher, 2005 [Publ 2682], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 10 10 1 --
  +----- Ambleminae      
--+                      
  |  +-- Unioninae       
  +--+                   
     +-- Margaritiferinae **

Amaro et al., 2017 [Publ 3791], fig. 3A-M

iTtTbiriSoTroTroSiD
1 1 0 1 6 6 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Konopleva et al., 2023 [Publ 4269], fig. S1_4

iTtTbiriSoTroTroSiD
1 1 0 1 55 55 1 --
  +-- Gibbosulinae **
--+               
  +-- Unionidae   

Lydeard et al., 2000 [Publ 2654], fig. 2d

iTtTbiriSoTroTroSiD
1 1 0 1 13 13 1 --
  +-- Unioninae       
  |                   
--+-- Margaritiferinae **
  |                   
  +-- Ambleminae      

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 17 12 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Gustafson & Iwamoto, 2005 [Publ 2741], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 4 4 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Araujo et al., 2009 [Publ 2589], fig. 7C

iTtTbiriSoTroTroSiD
1 1 0 1 9 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lydeard et al., 2000 [Publ 2654], fig. 2b

iTtTbiriSoTroTroSiD
1 1 0 1 13 13 1 --
  +-- Unioninae       
  |                   
--+-- Margaritiferinae **
  |                   
  +-- Ambleminae      

Prie et al., 2012 [Publ 3487], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 5 5 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 1 1 --
-- Margaritifera_margaritifera **

Zhang et al., 2013 [Publ 4214], fig. 5

iTtTbiriSoTroTroSiD
1 1 0 1 3 3 1 --
  +-------- Cristaria       
--+                         
  |  +----- Sinanodonta     
  +--+                      
     |  +-- Margaritiferinae **
     +--+                   
        +-- Gonideinae      

Araujo et al., 2009 [Publ 2589], fig. 7B

iTtTbiriSoTroTroSiD
1 1 0 1 10 10 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Hewitt et al., 2019 [Publ 3962], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 9 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 19 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Bogan et al., 2023 [Publ 4295], fig. 5

iTtTbiriSoTroTroSiD
1 1 0 1 5 5 1 --
  +-- Gibbosulinae **
--+               
  +-- Unionidae   

Reis et al., 2013 [Publ 3580], fig. 4

iTtTbiriSoTroTroSiD
1 1 0 1 13 12 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 4 1 1 --
-- Margaritifera_margaritifera **

Curole & Kocher, 2002 [Publ 2712], fig. 1-F

iTtTbiriSoTroTroSiD
1 1 0 1 3 3 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Bogan et al., 2023 [Publ 4295], fig. 4

iTtTbiriSoTroTroSiD
1 1 0 1 6 6 1 --
  +-- Gibbosulinae **
--+               
  +-- Unionidae   

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 1 1 --
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 1 1 --
-- Margaritifera_margaritifera **

Zhou et al., 2016 [Publ 3710], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 19 19 1 --
  +-------- Unioninae       
--+                         
  |  +----- Margaritiferinae **
  +--+                      
     |  +-- Ambleminae      
     +--+                   
        +-- Gonideinae      

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 6 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Turner et al., 2000 [Publ 2722], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 13 13 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 19 16 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Wu et al., 2020 [Publ 4147], fig. 3

iTtTbiriSoTroTroSiD
1 1 0 1 29 29 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 12 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 6 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lydeard et al., 2000 [Publ 2654], fig. 2a

iTtTbiriSoTroTroSiD
1 1 0 1 13 13 1 --
  +----- Unioninae       
  |                      
  +----- Margaritiferinae **
  |                      
--+----- Lampsilini      
  |                      
  +----- Amblemini       
  |                      
  |  +-- Quadrulini      
  +--+                   
     +-- Pleurobemini    

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 3 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Curole & Kocher, 2005 [Publ 2682], fig. 1-M

iTtTbiriSoTroTroSiD
1 1 0 1 11 11 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 8 1 --
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 3 1 --
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Froufe et al., 2017 [Publ 3802], fig. 7

iTtTbiriSoTroTroSiD
1 1 0 1 9 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Kano et al., 2019 [Publ 4194], fig. 6

iTtTbiriSoTroTroSiD
1 1 0 1 9 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Hoeh et al., 1998 [Publ 2695], fig. 3

iTtTbiriSoTroTroSiD
1 1 0 1 4 2 1 --
  +----- Unioninae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Ambleminae      

Amaro et al., 2017 [Publ 3791], fig. 3B-M

iTtTbiriSoTroTroSiD
1 1 0 1 6 6 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Zanatta & Murphy, 2006 [Publ 2672], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 55 55 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Araujo et al., 2017 [Publ 3801], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 13 13 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lydeard et al., 2000 [Publ 2654], fig. 2c

iTtTbiriSoTroTroSiD
1 1 0 1 13 13 1 --
  +----- Unioninae       
  |                      
  +----- Margaritiferinae **
  |                      
--+----- Quadrulini      
  |                      
  +----- Lampsilini      
  |                      
  |  +-- Amblemini       
  +--+                   
     +-- Pleurobemini    

Araujo et al., 2009 [Publ 2589], fig. 7A

iTtTbiriSoTroTroSiD
1 1 0 1 10 10 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Källersjö et al., 2005 [Publ 2684], fig. 1B-F

iTtTbiriSoTroTroSiD
1 1 0 1 6 6 1 --
  +-- Margaritiferinae **
  |                   
--+-- Anodontini      
  |                   
  +-- Unionini        

Inoue et al., 2014 [Publ 3635], fig. 3A

iTtTbiriSoTroTroSiD
1 1 0 1 19 19 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Lemer et al., 2019 [Publ 4093], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 5 1 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Froufe et al., 2019 [Publ 3936], fig. 4

iTtTbiriSoTroTroSiD
1 1 0 1 18 18 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Graf, 2000 [Publ 2647], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 16 6 1 --
-- Margaritifera **

Chen et al., 2020 [Publ 4170], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 33 33 1 --
                       +----- Prolasmidonta           
                    +--+                              
                    |  |  +-- Alasmidonta             
                    |  +--+                           
  +-----------------+     +-- Anemina                 
  |                 |                                 
  |                 |  +----- Pseudodontini           
  |                 +--+                              
  |                    |  +-- Lamprotula_caveata      
  |                    +--+                           
  |                       +-- Chamberlainia           
  |                                                   
  |                    +----- Oxynaiina               
  |              +-----+                              
  |              |     |  +-- Lepidodesmini           
--+              |     +--+                           
  |              |        +-- Aculamprotulina         
  |  +-----------+                                    
  |  |           |     +----- Platynaias              
  |  |           |  +--+                              
  |  |           |  |  |  +-- Utterbackia_imbecillis  
  |  |           |  |  +--+                           
  |  |           +--+     +-- Pyganodon               
  |  |              |                                 
  |  |              |  +----- Sinanodonta             
  |  |              +--+                              
  |  |                 |  +-- Cristaria               
  +--+                 +--+                           
     |                    +-- Anodontina              
     |                                                
     |                    +-- Utterbackia_peninsularis
     |              +-----+                           
     |              |     +-- Potamilus_alatus        
     |  +-----------+                                 
     |  |           |  +----- Lamprotula_leaii        
     |  |           +--+                              
     |  |              |  +-- Sinohyriopsis           
     +--+              +--+                           
        |                 +-- Gonideini               
        |                                             
        |  +----------------- Gibbosulinae **            
        +--+                                          
           |  +-------------- Quadrulini              
           +--+                                       
              |  +----------- Toxolasma               
              +--+                                    
                 |  +-------- Lampsilis_ornata        
                 +--+                                 
                    |  +----- Potamilus_leptodon      
                    +--+                              
                       |  +-- Lampsilis_siliquoidea   
                       +--+                           
                          +-- Lampsilis_powellii      

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 1 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Taylor et al., 2007 [Publ 2696], fig. 1-3

iTtTbiriSoTroTroSiD
1 1 0 1 3 2 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Froufe et al., 2019 [Publ 3936], fig. 3-M

iTtTbiriSoTroTroSiD
1 1 0 1 18 18 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 19 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Bogatov et al., 2018 [Publ 3859], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 5 5 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 1 1 --
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Campbell, 2000 [Publ 2724], fig. 1-5

iTtTbiriSoTroTroSiD
1 1 0 1 2 2 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 29 1 --
-- Margaritifera_margaritifera **

Källersjö et al., 2005 [Publ 2684], fig. 1A

iTtTbiriSoTroTroSiD
1 1 0 1 6 6 1 --
  +-- Margaritiferinae **
  |                   
--+-- Anodontini      
  |                   
  +-- Unionini        

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 9 1 --
-- Cumberlandia_monodonta **

Lopes-Lima et al., 2017 [Publ 3792], fig. 2-M

iTtTbiriSoTroTroSiD
1 1 0 1 9 9 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Hewitt et al., 2018 [Publ 3758], fig. 5

iTtTbiriSoTroTroSiD
1 1 0 1 8 8 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 12 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Pfeiffer et al., 2018 [Publ 3827], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 27 27 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 1 1 --
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Klishko et al., 2019 [Publ 3886], fig. 9

iTtTbiriSoTroTroSiD
1 1 0 1 10 10 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Huang et al., 2013 [Publ 3629], fig. 3

iTtTbiriSoTroTroSiD
1 1 0 1 15 15 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Bogatov et al., 2018 [Publ 3859], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 5 5 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Prié & Puillandre, 2014 [Publ 3529], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 16 16 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Reis et al., 2013 [Publ 3580], fig. 5

iTtTbiriSoTroTroSiD
1 1 0 1 14 13 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 3 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 12 1 --
  +-------------- Unioninae       
  |                               
--+  +----------- Lamprotulini    
  |  |                            
  +--+        +-- Gonideini       
     |  +-----+                   
     |  |     +-- Pseudodontini   
     +--+                         
        |  +----- Lampsilini      
        +--+                      
           |  +-- Pleurobemini    
           +--+                   
              +-- Margaritiferinae **

Giribet & Distel, 2003 [Publ 3670], fig. 3f5

iTtTbiriSoTroTroSiD
1 1 0 1 5 3 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 15 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae       

Zieritz et al., 2012 [Publ 3673], fig. 4

iTtTbiriSoTroTroSiD
1 1 0 1 7 7 1 --
  +----- Anodontina      
  |                      
--+  +-- Cristariina     
  |  |                   
  +--+-- Margaritiferinae **
     |                   
     +-- Unionini        

Graf et al., 2014 [Publ 3591], fig. 3

iTtTbiriSoTroTroSiD
1 1 0 1 10 10 1 --
  +-- Margaritiferinae **
  |                   
--+-- Unioninae       
  |                   
  +-- Parreysiinae    

Bogan et al., 2023 [Publ 4295], fig. 6

iTtTbiriSoTroTroSiD
1 1 0 1 12 12 1 --
  +-- Gibbosulinae **
--+               
  +-- Unionidae   

Araujo et al., 2018 [Publ 3779], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 17 17 1 --
  +-- Margaritiferinae **
--+                   
  +-- Unionidae